Song Sparrows Research Articles

Sparrow Preferences for Winter Cover Crops in California's Central Coast

Agriculture in many regions of the world has reduced bird habitat and abundance, and altered avian community structure. A study was conducted on an organic research farm over two winters (Oct to Mar) in an intensive agricultural region of Salinas Valley, CA to determine how cover crop variety and planting density influenced birds. Cover crops were rye (Secale cereale), a mixture of rye and legumes (Vicia spp., Pisum sativum), and a mustard mixture (Brassica juncea, Sinapis alba). White-crowned Sparrows (Zonotrichia leucophyrs), Savannah Sparrows (Passerculus sandwichensis) and Song Sparrows (Melospiza melodia) were observed both years in the study field. Bird droppings in cover crops were quantified and dissected to determine dietary preferences, and sparrow movement when flushed was determined. Dropping number and weight per m2 were at least 10 times greater in mustard than in rye and in the legume-rye mixture. Droppings were dominated by leaf tissue in mustard versus arthropod tissue in rye and legume-rye. Within cover crop variety, plant density did not have a clear or consistent effect on sparrows. Sparrows flushed from cover crops usually settled in mustard. The White-crowned Sparrow fed on mustard leaves and apparently on weed foliage under mustard. The arthropod-dominated droppings in rye and legume-rye cover crops were consistent with the food preferences of Song and Savannah Sparrows. The White-crowned Sparrow’s clear preference for mustard cover crops is likely due in part to their high dietary needs for sulfur-rich amino acids during the prenuptual molt. This paper provides novel information to help farmers and others understand the cover crop preferences of sparrows, and ways that farmers might use mustard cover crops as trap crops to reduce White-crowned Sparrow feeding damage on winter and spring vegetable crops. It also provides evidence of ecosystems services that these sparrows provide by feeding on weed tissue in winter cover crops.

Genomic differentiation and local adaptation on a microgeographic scale in a resident songbird.

Elucidating forces capable of driving species diversification in the face of gene flow remains a key goal in evolutionary biology. Song sparrows, Melospiza melodia, occur as 25 subspecies in diverse habitats across North America, are among the continent's most widespread vertebrate species, and are exemplary of many highly variable species for which the conservation of locally adapted populations may be critical to their range-wide persistence. We focus here on six morphologically distinct subspecies resident in the San Francisco Bay region, including three salt-marsh endemics and three residents in upland and riparian habitats adjacent to the Bay. We used reduced-representation sequencing to generate 2,773 SNPs to explore genetic differentiation, spatial population structure, and demographic history. Clustering separated individuals from each of the six subspecies, indicating subtle differentiation at microgeographic scales. Evidence of limited gene flow and low nucleotide diversity across all six subspecies further supports a hypothesis of isolation among locally adapted populations. We suggest that natural selection for genotypes adapted to salt marsh environments and changes in demography over the past century have acted in concert to drive the patterns of diversification reported here. Our results offer evidence of microgeographic specialization in a highly polytypic bird species long discussed as a model of sympatric speciation and rapid adaptation, and they support the hypothesis that conserving locally adapted populations may be critical to the range-wide persistence of similarly highly variable species.

No evidence that songbirds use odour cues to avoid malaria-infected conspecifics

Abstract Many animals have evolved mechanisms to detect and avoid parasitized conspecifics, primarily through odour cues, but whether birds are capable of odour-mediated parasite avoidance is unknown. Recently, we showed that exposing song sparrows (Melospiza melodia) to avian malaria parasites (Plasmodium sp.) alters the chemical composition of their preen oil, which is the major source of body odour in birds. Here, we presented song sparrows with preen oil from uninfected (sham-inoculated) and malaria-infected conspecifics, predicting that birds would spend more time with odour cues from uninfected than infected birds. Birds without detectable malarial infections spent about 50% more time with preen oil from uninfected than infected conspecifics, and females spent nearly twice as much time with preen oil from uninfected than infected conspecifics. However, neither difference was statistically significant. Song sparrows may be able to detect odour cues of infection, but further experiments are needed to confirm or refute this.

İran’da Toplumsal Gerçeklik Ve Sinema İlişkisi Üzerine Dönemsel Bir Karşılaştırma: Sohrab Şehide Sales’in “Cansız Tabiat” Ve Mecid Mecidi’nin “Serçelerin Şarkısı”

Sinema sanatı, diğer ülkelerde olduğu gibi İran’da da siyasal, toplumsal ve kültürel değişimlerden etkilenmiş ve bu doğrultuda şekillenmiştir. Doğası gereği içinde bulunduğu dönemin şartlarına tanıklık eden sinema; İran’ın sosyal, siyasal, ekonomik ve kültürel yapısını geniş kitlelere ulaştırır. Bu bağlamda sinema filmleri; belli bir dönemin toplumsal ve siyasal yapısını kavrayabilmek ve söz konusu döneme dair çıkarımlarda bulunabilmek adına, referans olarak başvurulabilecek üretimlerdir. Bu çalışmada; İran’da toplumsal yaşam kadar kültürel üretim açısından da bir dönüm noktası olan İslam Devrimi’nin gerçekleştiği 1979 yılı öncesi ve sonrası döneminin sosyal ve siyasal yapısındaki benzerlikler ve farklılıkların sinema üzerinden incelenmesi amaçlanmıştır. Bu amaç doğrultusunda; İran sinemasının önemli temsilcilerinden Sohrab Şehide Sales’in “Cansız Tabiat” (1975) adlı filmi ile Mecid Mecidi’nin “Serçelerin Şarkısı” (2008) filmi incelemeye esas alınmıştır. Üretimlerini farklı dönemlerde gerçekleşmiş bu iki yönetmen de; filmlerinde dönemlerinin toplumsal gerçeklerini, İran’ın geleneksel aile yapısını, sosyal yaşantısını, kadın sorunlarını, siyasi sorunları, yoksulluğu ve işsizlik problemini konu edinmiştir. Bu bakımdan “Cansız Tabiat” ve “Serçelerin Şarkısı” ele aldıkları konular üzerinden bize dönemin toplumsal, siyasal ve ekonomik yapısının analizine imkân veren filmlerdir. Çalışma kapsamında, filmler, arasındaki benzerlikler ve farklılıklar bakımından karşılaştırılacak ve bu doğrultuda, her iki filmin çekildiği dönemin toplumsal ve siyasal yapısına yönelik bir değerlendirilme yapılacaktır.

Social factors in bird-song development: Learning to sing with friends and rivals.

Laboratory studies have revealed that social factors are key in bird-song learning. Nevertheless, little is known about how or why birds choose the songs they do learn from the many they will hear under natural conditions. We focus on various theories concerning social song learning that have been offered to date, with special attention paid to two axes of social factors. First, does song learning occur via direct interaction of the young bird with song tutors, or via social eavesdropping by the young bird on interacting singers (social modeling of song)? Social modeling, a hypothesis first proposed by Pepperberg (Zeitschrift für Tierpsychologie, 55(2), 139-160, 1981), and direct interaction are not mutually exclusive hypotheses, and the evidence we review suggests both play a role in song learning. Second, does song learning occur via interactions with rivals (territorial competitors) or with friends (mutually tolerant or even cooperative territorial neighbors). These are largely mutually exclusive hypotheses, and can really only be tested in the field. There is little evidence on this contrast to date. We review our recent study on song sparrows, which indicates that both the young bird and his primary tutor may benefit from song learning/tutoring. If this mutual benefit result is confirmed by further studies, we believe that song "tutoring" in these cases may be more than a term of convenience: that it may qualify as true teaching.

Animal Culture: How a New Birdsong Went Viral

A new white-throated sparrow song has overtaken most of Canada in less than 20 years. The explanation for this remarkably fast spread may lie in the southern migratory grounds, where populations from across Canada converge each winter.

Birdsong learning is mutually beneficial for tutee and tutor in song sparrows

Song learning is generally assumed to be beneficial for a young songbird, but merely incidental, without costs or benefits, for the older song ‘tutors’. In the present study we contrast two mutually exclusive hypotheses about the tutor/tutee relationship: (1) that it is cooperative, or at least mutually tolerant, with tutor and tutee mutually benefiting from their relationship, versus (2) that it is competitive, with tutor and tutee competing over territory, so that one or the other suffers negative fitness consequences of their relationship. In a field study of three consecutive cohorts of song sparrows, Melospiza melodia morphna, we determined the older bird (primary tutor) from whom the young bird (tutee) learned most of his songs, and how long tutee and primary tutor survived subsequently. We found that the more songs a tutee learns from his primary tutor, the longer their mutual survival on their respective territories. While the number of songs they share predicts the mutual survival of tutor and tutee, it does not predict the independent survival of tutor or tutee, suggesting that the benefit each receives from song sharing exists only so long as both survive.

Song sparrows do not discriminate between their own song and stranger song

Bird song is socially learned. During song learning, the bird’s hearing its own vocalization is important for normal development of song. Whether bird’s own song is represented and recognized as a special category in adult birds, however, is unclear. If birds respond differently to their own songs when these are played back to them, this would be evidence for auditory self-recognition. To test this possibility, we presented song sparrow males (Melospiza melodia) playbacks of their own songs or stranger songs and measured aggressive responses as well as type matching. We found no evidence of behavioral discrimination of bird’s own song relative to the (non-matching) stranger song. These findings cast doubt on an earlier proposal that song sparrows display auditory self-recognition and support the common assumption in playback experiments that bird’s own song is perceived as stranger song.

Effects of low-density urbanization on genetic structure in the Song Sparrow.

Urbanization fragments landscapes and can impede the movement of organisms through their environment, which can decrease population connectivity. Reduction in connectivity influences gene flow and allele frequencies, and can lead to a reduction in genetic diversity and the fixation of certain alleles, with potential negative effects for populations. Previous studies have detected effects of urbanization on genetic diversity and structure in terrestrial animals living in landscapes that vary in their degree of urbanization, even over very short distances. We investigated the effects of low-intensity urbanization on genetic diversity and genetic structure in Song Sparrows (Melospiza melodia). We captured 208 Song Sparrows at seven sites along a gradient of urbanization in and around Blacksburg, VA, USA, then genotyped them using a panel of fifteen polymorphic microsatellite loci. We found that genetic diversity was comparable among the seven study sites, and there was no evidence of genetic structuring among sites. These findings suggest that over a gradient of urbanization characterized by low density urban development, Song Sparrows likely exist in a single panmictic population.

Bird Species Involved in West Nile Virus Epidemiological Cycle in Southern Québec.

Despite many studies on West Nile Virus (WNV) in the US, including the reservoir role of bird species and the summer shifts of the Culex mosquito, feeding from birds to mammals, there have been few equivalent studies in the neighboring regions of Canada where WNV is endemic. Here, a priority list of bird species likely involved in WNV transmission in the greater Montréal area is constructed by combining three sources of data: (i) from WNV surveillance in wild birds (2002–2015); (ii) blood meal analysis of Culex pipiens–restuans (CPR), the primary enzootic vectors of WNV in the region, collected from surveillance in 2008 and 2014; (iii) literature review on the sero-prevalence/host competence of resident birds. Each of these data sources yielded 18, 23 and 53 species, and overall, 67 different bird species were identified as potential WNV amplifiers/reservoirs. Of those identified from CPR blood meals, Common starlings, American robins, Song sparrows and House sparrows ranked the highest and blood meal analysis demonstrated a seasonal shift in feed preference from birds to mammals by CPR. Our study indicates that there are broad similarities in the ecology of WNV between our region and the northeastern US, although the relative importance of bird species varies somewhat between regions.

A High-Quality Genome Assembly of the North American Song Sparrow, Melospiza melodia.

The song sparrow, Melospiza melodia, is one of the most widely distributed species of songbirds found in North America. It has been used in a wide range of behavioral and ecological studies. This species’ pronounced morphological and behavioral diversity across populations makes it a favorable candidate in several areas of biomedical research. We have generated a high-quality de novo genome assembly of M. melodia using Illumina short read sequences from genomic and in vitro proximity-ligation libraries. The assembled genome is 978.3 Mb, with a physical coverage of 24.9×, N50 scaffold size of 5.6 Mb and N50 contig size of 31.7 Kb. Our genome assembly is highly complete, with 87.5% full-length genes present out of a set of 4,915 universal single-copy orthologs present in most avian genomes. We annotated our genome assembly and constructed 15,086 gene models, a majority of which have high homology to related birds, Taeniopygia guttata and Junco hyemalis. In total, 83% of the annotated genes are assigned with putative functions. Furthermore, only ∼7% of the genome is found to be repetitive; these regions and other non-coding functional regions are also identified. The high-quality M. melodia genome assembly and annotations we report will serve as a valuable resource for facilitating studies on genome structure and evolution that can contribute to biomedical research and serve as a reference in population genomic and comparative genomic studies of closely related species.

Recent immigrants alter the quantitative genetic architecture of paternity in song sparrows.

Quantifying additive genetic variances and cross‐sex covariances in reproductive traits, and identifying processes that shape and maintain such (co)variances, is central to understanding the evolutionary dynamics of reproductive systems. Gene flow resulting from among‐population dispersal could substantially alter additive genetic variances and covariances in key traits in recipient populations, thereby altering forms of sexual conflict, indirect selection, and evolutionary responses. However, the degree to which genes imported by immigrants do in fact affect quantitative genetic architectures of key reproductive traits and outcomes is rarely explicitly quantified. We applied structured quantitative genetic analyses to multiyear pedigree, pairing, and paternity data from free‐living song sparrows (Melospiza melodia) to quantify the differences in mean breeding values for major sex‐specific reproductive traits, specifically female extra‐pair reproduction and male paternity loss, between recent immigrants and the previously existing population. We thereby quantify effects of natural immigration on the means, variances, and cross‐sex covariance in total additive genetic values for extra‐pair paternity arising within the complex socially monogamous but genetically polygynandrous reproductive system. Recent immigrants had lower mean breeding values for male paternity loss, and somewhat lower values for female extra‐pair reproduction, than the local recipient population, and would therefore increase the emerging degree of reproductive fidelity of social pairings. Furthermore, immigration increased the variances in total additive genetic values for these traits, but decreased the magnitudes of the negative cross‐sex genetic covariation and correlation below those evident in the existing population. Immigration thereby increased the total additive genetic variance but could decrease the magnitude of indirect selection acting on sex‐specific contributions to paternity outcomes. These results demonstrate that dispersal and resulting immigration and gene flow can substantially affect quantitative genetic architectures of complex local reproductive systems, implying that comprehensive theoretical and empirical efforts to understand mating system dynamics will need to incorporate spatial population processes.

Birds suppress pests in corn but release them in soybean crops within a mixed prairie/agriculture system.

Birds provide ecosystem services (pest control) in many agroecosystems and have neutral or negative ecological effects (disservices) in others. Large-scale, conventional row crop agriculture is extremely widespread globally, yet few studies of bird effects take place in these agroecosystems. We studied indirect effects of insectivorous birds on corn and soybean crops in fields adjacent to a prairie in Illinois (USA). We hypothesized that prairie birds would forage for arthropods in adjacent crop fields and that the magnitude of services or disservices would decrease with distance from the prairie. We used bird-excluding cages over crops to examine the net effect of birds on corn and soybean grain yield. We also conducted DNA metabarcoding to identify arthropod prey in fecal samples from captured birds. Our exclosure experiments revealed that birds provided net services in corn and net disservices in soybeans. Distance from prairie was not a significant predictor of exclosure treatment effect in either crop. Many bird fecal samples contained DNA from both beneficial arthropods and known economically significant pests of corn, but few economically significant pests of soybeans. Song Sparrows (Melospiza melodia), one of our most captured species, most commonly consumed corn rootworms, an economically significant pest of corn crops. We estimated that birds in this system provided a service worth approximately US $275 ha-1 in corn yield gain, and a disservice valued at approximately $348 ha-1 in soybean yield loss. Our study is the first to demonstrate that birds can provide substantial and economically valuable services in field corn, and disservices in soybean crops. The contrasting findings in the 2 crop systems suggest a range of bird impacts within widespread agroecosystems and demonstrate the importance of quantifying net trophic effects.

Territory establishment, song learning strategies and survival in song sparrows

AbstractIn most songbirds, the processes of song learning and territory establishment overlap in the early life and a young bird usually winds up with songs matching those of his territorial neighbors in his first breeding season. In the present study, we examined the relationships among the timing of territory establishment, the pattern of song learning and territorial success in a sedentary population of song sparrows (Melospiza melodia). Males in this population tend to learn their songs from their neighbors and consequently they show high song sharing with neighbors and use these shared songs preferentially in interactions with them. Males also show significant variation in the timing of territory establishment, ranging from their natal summer to the next spring. Using a three‐year dataset, we found that the timing of territory establishment did not systematically affect the composition of the song repertoire of the tutee: early establishers and late establishers learned equally as much from their primary tutors and had a similar number of tutors and similar repertoire sizes, nor did timing of territory establishment affect subsequent survival on territory. Therefore, the song‐learning program of song sparrows seems versatile enough to lead to high song sharing even for birds that establish territories relatively late.

Food stress, but not experimental exposure to mercury, affects songbird preen oil composition.

Mercury is a global pollutant and potent neurotoxic metal. Its most toxic and bioavailable form, methylmercury, can have both lethal and sublethal effects on wildlife. In birds, methylmercury exposure can disrupt behavior, hormones, the neuroendocrine system, and feather integrity. Lipid-rich tissues and secretions may be particularly susceptible to disruption by lipophilic contaminants such as methylmercury. One such substance is feather preen oil, a waxy secretion of the uropygial gland that serves multiple functions including feather maintenance, anti-parasitic defense, and chemical signaling. If methylmercury exposure alters preen oil composition, it could have cascading effects on feather quality, susceptibility to ectoparasites, and mate choice and other social behaviors. We investigated whether exposure to methylmercury, either alone or in association with other stressors, affects preen oil chemical composition. We used a two-factor design to expose adult song sparrows (Melospiza melodia) to an environmentally relevant dietary dose of methylmercury and/or to another stressor (unpredictable food supply) for eight weeks. The wax ester composition of preen oil changed significantly over the 8-week experimental period. This change was more pronounced in the unpredictable food treatment, regardless of dietary methylmercury. Contrary to our prediction, we found no main effect of methylmercury exposure on preen oil composition, nor did methylmercury interact with unpredictable food supply in predicting the magnitude of chemical shifts in preen oil. While it remains critical to study sublethal effects of methylmercury on wildlife, our findings suggest that the wax ester composition of preen oil is robust to environmentally relevant doses of this contaminant.

Brain activity underlying American crow processing of encounters with dead conspecifics

Animals utilize a variety of auditory and visual cues to navigate the landscape of fear. For some species, including corvids, dead conspecifics appear to act as one such visual cue of danger, and prompt alarm calling by attending conspecifics. Which brain regions mediate responses to dead conspecifics, and how this compares to other threats, has so far only been speculative. Using 18F-fluorodeoxyglucose positron emission tomography (FDG-PET) we contrast the metabolic response to visual and auditory cues associated with a dead conspecific among five a priori selected regions in the American crow (Corvus brachyrhynchos) brain: the hippocampus, nidopallium caudolaterale, striatum, amygdala, and the septum. Using a repeated-measures, fully balanced approach, we exposed crows to four stimuli: a dead conspecific, a dead song sparrow (Melospiza melodia), conspecific alarm calls given in response to a dead crow, and conspecific food begging calls. We find that in response to observations of a dead crow, crows show significant activity in areas associated with higher-order decision-making (NCL), but not in areas associated with social behaviors or fear learning. We do not find strong differences in activation between hearing alarm calls and food begging calls; both activate the NCL. Lastly, repeated exposures to negative stimuli had a marginal effect on later increasing the subjects’ brain activity in response to control stimuli, suggesting that crows might quickly learn from negative experiences.

Development of the corticosterone stress response differs among passerine species

Glucocorticoids are steroid hormones which increase dramatically in response to a physical or perceived stressor. However, developing young of altricial species typically have a damped glucocorticoid stress response. The developmental hypothesis posits that the physiological stress response should develop concurrently with an individual’s ability to respond to a challenge. The dampened response may benefit an organism, as chronic exposure to glucocorticoids can have short- and long-term detrimental effects, and altricial young are unable to escape most stressors. However, we do not know if or why species with similar ontogeny vary in their development of the physiological stress response. We assessed levels of baseline and stress-induced corticosterone (the main avian glucocorticoid) in six passerine species with varying life-history strategies, including a brood parasite, the brown-headed cowbird (Molothrus ater). Circulating baseline corticosterone levels increased with nestling age for all species. Stress-induced corticosterone levels sampled at 15-min post-capture significantly increased with nestling age at a similar rate and magnitude in brown-headed cowbirds, eastern phoebes (Sayornis phoebe), hooded warblers (Setophaga citrina), red-winged blackbirds (Agelaius phoeniceus), and song sparrows (Melospiza melodia). However, gray catbird (Dumetella carolinensis) nestlings showed an extremely dampened elevation in corticosterone in response to stress, even near fledge. Gray catbirds are unusual among songbirds, as they are open-ended song learners. Stress during development can profoundly influence avian song learning and performance abilities. However, further study is necessary to determine if there is a relationship between nestling adrenocortical activity and open- versus closed-ended song learning.

Impacts of Riparian Restoration on Vegetation and Avifauna on Private and Communal Lands in Northwest Mexico and Implications for Future Efforts

Restoring and enhancing riparian vegetation on private and communal lands in Mexico is important for biodiversity conservation given the ecological significance of these areas and the scarcity of public protected areas. To enhance riparian vegetation and wildlife habitats and train local people in restoration techniques, we implemented restoration and outreach efforts on private and communal lands in the Sky Islands region of northwest Mexico. We fenced 475 ha of riparian zones from livestock, erected erosion-control structures, planted trees, and developed management agreements for cool-season grazing with landowners on 10 ranches across 3 sites in 2012-2013, then repaired fences and renegotiated agreements in 2017-2019. To foster evaluation, we used a before-after/control-impact design to measure attributes of vegetation structure and bird communities and compared baselines from 2012 with post-treatment estimates from 2019. As predicted, understory vegetation volume generally increased in treatments relative to controls ( P = .09), especially when one treatment area with the lowest pre-treatment grazing impacts was censored ( P = .01). Although canopy cover also increased, there was little differential change in treatments relative to controls ( P ⩾ .23) due likely to longer time periods needed to realize responses. Densities of most focal bird populations varied across time periods in directions that typically matched observed changes in vegetation structure, but fewer species showed signs of differential positive change linked to treatments relative to controls. Densities of Yellow-breasted Chat, a key understory obligate and important focal species, increased in treatments relative to controls across sites, as did densities of Sinaloa Wren, which also use dense underbrush ( P ⩽ .05). Positive changes by other understory obligates (eg, Common Yellowthroat, Song Sparrow) were more local but sometimes of high magnitude (>8-fold) also suggesting positive impacts of treatments. Despite mixed results over a limited time period, these patterns suggest restoration efforts drove localized recovery of understory vegetation and associated bird populations, but benefits varied widely with environmental and social factors linked to management. Greater ecological benefits to riparian areas on private and communal lands in this region can be fostered by further incentivizing construction, maintenance, and proper use of restoration infrastructure, through education, and by building relationships based on trust and credibility with landowners.

Perceived wintering latitude determines timing of song output in a migratory bird.

Migratory bird populations frequently consist of individuals that overwinter variable distances from the breeding site. Seasonal changes in photoperiod, which varies with latitude, underlie seasonal changes in singing frequency in birds. Therefore, migratory populations that consist of individuals that overwinter at different latitudes with large overwintering ranges could experience within‐population variation in seasonal production of song. To test the influence of overwintering latitude on intrapopulation variance in song production in the spring, we subjected two groups of Eastern Song Sparrows (Melospiza melodia melodia) from the same partially migratory breeding population to different photoperiodic schedules associated with a 1,300‐km difference in overwintering location. One group remained on the natural photoperiodic schedule of the breeding site (resident group) while the other group experienced a nonbreeding photoperiod that mimicked a southern migration in the fall followed by a northern migration back to the breeding site in the spring (migratory group). We compared song output between the two groups in three different stages (nonbreeding, prebreeding, and breeding). Little singing occurred during nonbreeding stage sample dates (20 November, 6 December) for the resident group, and no singing occurred for the migrant group. During the prebreeding stage (27 January, 7 February), significantly more singing occurred in the resident group than in the migrant group. During the breeding stage (21 March, 4 April), after a simulated migration for the migrants, song output was similar in both groups. These results suggest that within‐population variation in wintering latitude may contribute to variation in seasonal changes in singing behavior, which may covary with readiness to breed. Studies utilizing confirmed migrants and residents, rather than merely simulated migrants and residents, are also needed to better understand these processes.

Testing predictions of inclusive fitness theory in inbreeding relatives with biparental care.

Inclusive fitness theory predicts that parental care will vary with relatedness between potentially caring parents and offspring, potentially shaping mating system evolution. Systems with extra-pair paternity (EPP), and hence variable parent–brood relatedness, provide valuable opportunities to test this prediction. However, existing theoretical and empirical studies assume that a focal male is either an offspring's father with no inbreeding, or is completely unrelated. We highlight that this simple dichotomy does not hold given reproductive interactions among relatives, complicating the effect of EPP on parent–brood relatedness yet providing new opportunities to test inclusive fitness theory. Accordingly, we tested hierarchical hypotheses relating parental feeding rate to parent–brood relatedness, parent kinship and inbreeding, using song sparrows (Melospiza melodia) experiencing natural variation in relatedness. As predicted, male and female feeding rates increased with relatedness to a dependent brood, even controlling for brood size. Male feeding rate tended to decrease as paternity loss increased, and increased with increasing kinship and hence inbreeding between socially paired mates. We thereby demonstrate that variation in a key component of parental care concurs with subtle predictions from inclusive fitness theory. We additionally highlight that such effects can depend on the underlying social mating system, potentially generating status-specific costs of extra-pair reproduction.