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The songs of birds are complex signals that may have several functions and vary widely among species. Different ecological, behavioural and morphological factors, as well as phylogeny, have been associated as predictors of the evolution of song structure. However, the importance of differences in development, despite their relevance, has seldom been considered. Here, we analysed the evolution of song in two families of songbirds that differ in song development, manakins (suboscines) and cardinals (oscines), with their phylogeny, morphology, and ecology. Our results show that song characteristics had higher phylogenetic signal in cardinals than in manakins, suggesting higher evolutionary lability in the suboscines. Body mass was the main predictor of song parameters in manakins, and together with habitat type, had a major effect on cardinals' song structure. Precipitation and altitude were also associated with some song characteristics in cardinals. Our results bring unexpected insights into birdsong evolution, in which non-learners (manakins) revealed greater evolutionary lability than song learners (cardinals)., (© 2024. The Author(s).)

In most avian retinas, double cones (consisting of a principal and accessory member) outnumber other photoreceptor types and have been associated with various functions, such as encoding luminance, sensing polarized light, and magnetoreception. However, their down-stream circuitry is poorly understood, particularly across bird species. Analysing species differences is important to understand changes in circuitry driven by ecological adaptations. We compare the ultrastructure of double cones and their postsynaptic bipolar cells between a night-migratory European robin and non-migratory chicken. We discover four previously unidentified bipolar cell types in the European robin retina, including midget-like bipolar cells mainly connected to one principal member. A downstream ganglion cell reveals a complete midget-like circuit similar to a circuit in the peripheral primate retina. Additionally, we identify a selective circuit transmitting information from a specific subset of accessory members. Our data highlight species-specific differences in double cone to bipolar cell connectivity, potentially reflecting ecological adaptations., (© 2024. The Author(s).)

Sexual signals such as colour ornamentation and birdsong evolve independently of each other in some clades, and in others they evolve positively or negatively correlated. We rarely know why correlated evolution does or does not occur. Here, we show positively correlated evolution between plumage colour and song motor performance among canaries, goldfinches and allies, associated with species differences in body size. When controlling for body size, the pattern of correlated evolution between song performance and colour disappeared. Syllable diversity was not as strongly associated with size, and did not evolve in a correlated manner with colour. We argue that correlated evolution between song and colour was mediated by large size limiting song motor performance, likely due to constraints on the speed of moving heavier bills, and by larger species having less saturated plumage colour, possibly due to life-history traits of larger birds (e.g. longevity, stable pairs) contributing to weaker sexual selection. Results are consistent with the hypothesis that correlated evolution between sexual signals is influenced by how, in a clade, selective pressures and constraints affecting each type of signal happen to be co-distributed across species. Such contingency helps explain the diversity in clade-specific patterns of correlated evolution between sexual signals.

The magnetic compass sense of migratory songbirds is thought to derive from magnetically sensitive photochemical reactions in cryptochromes located in photoreceptor cells in the birds' retinas. More specifically, transient radical pairs formed by light-activation of these proteins have been proposed to account for the birds' ability to orient themselves using the Earth's magnetic field and for the observation that radiofrequency magnetic fields, superimposed on the Earth's magnetic field, can disrupt this ability. Here, by means of spin dynamics simulations, we show that it may be possible for the birds to orient in a monochromatic radiofrequency field in the absence of the Earth's magnetic field. If such a behavioural test were successful, it would provide powerful additional evidence for a radical pair mechanism of avian magnetoreception.

During spring migration, nocturnal migrants attempt to minimize their travel time to reach their breeding grounds early. However, how they behave and respond to unfavourable conditions during their springtime travels is much less understood. In this study, we reveal the effects of atmospheric factors on nocturnal bird migration under adverse conditions during spring and autumn, based on one of the most detailed bird migration studies globally, using radar data from 13 deployments over a period of seven years (2014-2020) in the Levant region. Using ERA5 reanalysis data, we found that migratory birds maintain similar ground speeds in both autumn and spring migrations, but during spring, when encountering unfavourable winds, they put more effort into maintaining their travel speed by increasing self-powered airspeed by 18%. Moreover, we report for the first time that spring migrants showed less selectivity to wind conditions and migrated even under unfavourable headwind and crosswind conditions. Interestingly, we discovered that temperature was the most important weather parameter, such that warm weather substantially increased migration intensities in both seasons. Our results enhance our understanding of bird migration over the Levant region, one of the world's largest and most important migration flyways, and the factors controlling it. This information is essential for predicting bird migration, which-especially under the ongoing anthropogenic changes-is of high importance.

Vocalization is an important communication tool that can reflect many aspects of an individual's internal and external condition. This is especially true for birds. Previous research has shown that bird calls and songs change in response to a variety of potential stressors, although the extent and direction of the changes depend on the nature of the stressor and the environment. Circulating glucocorticoids, such as corticosterone, often increase in response to stressors and mediate some of the observed changes via alterations of the individual's physiological state. Acute elevations of corticosterone often occur as a physiological response to short-term stressors; however, the effects of this elevation on adult vocalizations have not been well documented. Here, we experimentally elevated corticosterone at two different levels using a noninvasive method and examined the effects on the vocal communication of male and female adult common mynas (Acridotheres tristis). Corticosterone elevation temporarily increased song output and some measures of song complexity, while call output decreased. These effects were dosage dependent (higher corticosterone levels had a stronger effect), most evident 40 min after ingestion, and some vocal changes were sex-specific. Future studies should investigate whether the changes in vocal performance due to elevated glucocorticoids have consequences for the birds' behavior, reproductive success, and survival., (© 2024 The Authors. Journal of Experimental Zoology Part A: Ecological and Integrative Physiology published by Wiley Periodicals LLC.)

Migrating animals perform astonishing seasonal movements by orienting and navigating over thousands of kilometres with great precision. Many migratory species use cues from the sun, stars, landmarks, olfaction and the Earth's magnetic field for this task. Among vertebrates, songbirds are the most studied taxon in magnetic-cue-related research. Despite multiple studies, we still lack a clear understanding of when, where and how magnetic cues affect the decision-making process of birds and hence, their realised migratory behaviour in the wild. This understanding is especially important to interpret the results of laboratory experiments in an ecologically appropriate way. In this review, we summarise the current findings about the role of magnetic cues for migratory decisions in songbirds. First, we review the methodological principles for orientation and navigation research, specifically by comparing experiments on caged birds with experiments on free-flying birds. While cage experiments can show the sensory abilities of birds, studies with free-flying birds can characterise the ecological roles of magnetic cues. Second, we review the migratory stages, from stopover to endurance flight, in which songbirds use magnetic cues for their migratory decisions and incorporate this into a novel conceptual framework. While we lack studies examining whether and when magnetic cues affect orientation or navigation decisions during flight, the role of magnetic cues during stopover is relatively well studied, but mostly in the laboratory. Notably, many such studies have produced contradictory results so that understanding the biological importance of magnetic cues for decisions in free-flying songbirds is not straightforward. One potential explanation is that reproducibility of magnetic-cue experiments is low, probably because variability in the behavioural responses of birds among experiments is high. We are convinced that parts of this variability can be explained by species-specific and context-dependent reactions of birds to the study conditions and by the bird's high flexibility in whether they include magnetic cues in a decision or not. Ultimately, this review should help researchers in the challenging field of magnetoreception to design experiments meticulously and interpret results of such studies carefully by considering the migration ecology of their focal species., (© 2024 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

The distribution of a species reflects its ecological adaptability and evolutionary history, which is shaped by the environment and represents a dynamic area subject to anthropogenic environmental change. We used the MaxEnt algorithm to construct ecological niche models for four thrush species within the Turdus genus; T. amaurochalinus, T. chiguanco, T. falcklandii and T. rufiventris. These models were used to predict the potential geographic distributions of these species that are expanding their ranges in South America. Using occurrence records, we estimated currently occupied areas for each species. We also identified suitable habitats and projected possible areas to be colonized by the four species at continental scale. Temperature annual range had the highest influence for T. falcklandii, while human modification was the main variable explaining the distribution of the other three species. The potential distribution area ranged from 2.5 million km 2 for T. falcklandii to nearly seven million km 2 for T. amaurochalinus. Large proportions of suitable area remain unoccupied by all four species, being 50% for T. amaurochalinus and T. rufiventris, and about 70% for T. chiguanco and T. falcklandii. Anthropogenic disturbances, such as habitat loss and ecosystem transformation, lead to non-random species extinction and biotic homogenization, highlighting the importance of predictive models as valuable tools for informing mitigation policies and conservation strategies. Thrushes are progressively expanding their ranges, and the colonization of new habitats could bring new challenges., (© 2024. The Author(s).)

It has recently become clear that some language-specific traits previously thought to be unique to humans (such as the capacity to combine sounds) are widespread in the animal kingdom. Despite the increase in studies documenting the presence of call combinations in non-human animals, factors promoting this vocal trait are unclear. One leading hypothesis proposes that communicative complexity co-evolved with social complexity owing to the need to transmit a diversity of information to a wider range of social partners. The Western Australian magpie ( Gymnorhina tibicen dorsalis ) provides a unique model to investigate this proposed link because it is a group-living, vocal learning species that is capable of multi-level combinatoriality (independently produced calls contain vocal segments and comprise combinations). Here, we compare variations in the production of call combinations across magpie groups ranging in size from 2 to 11 birds. We found that callers in larger groups give call combinations: (i) in greater diversity and (ii) more frequently than callers in smaller groups. Significantly, these observations support the hypothesis that combinatorial complexity may be related to social complexity in an open-ended vocal learner, providing an important step in understanding the role that sociality may have played in the development of vocal combinatorial complexity. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'.

Birds maintain some of the highest body temperatures among endothermic animals. Often deemed a selective advantage for heat tolerance, high body temperatures also limits birds' thermal safety margin before reaching lethal levels. Recent modelling suggests that sustained effort in Arctic birds might be restricted at mild air temperatures, which may require reductions in activity to avoid overheating, with expected negative impacts on reproductive performance. We measured within-individual changes in body temperature in calm birds and then in response to an experimental increase in activity in an outdoor captive population of Arctic, cold-specialised snow buntings (Plectrophenax nivalis), exposed to naturally varying air temperatures (- 15 to 36 °C). Calm buntings exhibited a modal body temperature range from 39.9 to 42.6 °C. However, we detected a significant increase in body temperature within minutes of shifting calm birds to active flight, with strong evidence for a positive effect of air temperature on body temperature (slope = 0.04 °C/ °C). Importantly, by an ambient temperature of 9 °C, flying buntings were already generating body temperatures ≥ 45 °C, approaching the upper thermal limits of organismal performance (45-47 °C). With known limited evaporative heat dissipation capacities in these birds, our results support the recent prediction that free-living buntings operating at maximal sustainable rates will increasingly need to rely on behavioural thermoregulatory strategies to regulate body temperature, to the detriment of nestling growth and survival., (© 2024. The Author(s).)

Phenological adjustment is the first line of adaptive response of vertebrates when seasonality is disrupted by climate change. The prevailing response is to reproduce earlier in warmer springs, but habitat changes, such as forest degradation, are expected to affect phenological plasticity, for example, due to loss of reliability of environmental cues used by organisms to time reproduction. Relying on a two-decade, country-level capture-based monitoring of common songbirds' reproduction, we investigated how habitat anthropization, here characterized by the rural-urban and forest-farmland gradients, affected the average phenology and plasticity to local temperature in two common species, the great tit Parus major and the blue tit Cyanistes caeruleus. We built a hierarchical model that simultaneously estimated fledging phenology and its response to spring temperatures based on the changes in the proportion of juveniles captured over the breeding season. Both species fledge earlier in warmer sites (blue tit: 2.94 days/°C, great tit: 3.83 days/°C), in warmer springs (blue tit: 2.49 days/°C, great tit: 2.75 days/°C) and in most urbanized habitats (4 days for blue tit and 2 days for great tit). The slope of the reaction norm of fledging phenology to spring temperature varied across sites in both species, but this variation was explained by habitat anthropization only in the deciduous forest specialist, the blue tit. In this species, the responses to spring temperature were shallower in agricultural landscapes and slightly steeper in more urban areas. Habitat anthropization did not explain variation in the slope of the reaction norm in the habitat-generalist species (great tit), for which mean fledgling phenology and plasticity were correlated (i.e., steeper response in later sites). The effects of habitat change on phenological reaction norms provide another way through which combined environmental degradations may threaten populations' persistence, to an extent depending on species and on the changes in their prey phenology and abundance., (© 2024 The Author(s). Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Climate change is altering the timing of seasonal events for many taxa. There is limited understanding of how northward/southward songbird migration follows or is limited by the latitudinal progression of seasonal transitions. Consistent environmental conditions that migrating birds encounter across latitudes likely represent or correlate with important resources or limiting factors for migration. We tested whether migratory passage-observed via radar-consistently tracked land surface variables and phenophases across latitudes in the US Central Flyway in both spring and fall. The daily temperatures, precipitation and vegetation greenness occurring on 10%, 50% and 90% cumulative passage dates changed substantially with latitude, indicating that most migrants experienced rapidly changing conditions as they headed north or south. Temperature did not limit the progression of migration in either season. Peak spring migration in the southern US occurred nearly 40 days after the spring green wave, the northward progression of vegetation growth, but nearly caught up to green-up at 48° N. Spring migration phenology may have evolved to prioritize earlier arrival for breeding. Across all latitudes, peak fall migration coincided with the same land surface phenophase, an interval of 26 days prior to dormancy onset. Migrants may rely on phenological events in vegetation during fall stopovers. Considering that (a) migratory passage tracked fall land surface phenology across latitudes at a continental scale, (b) previous studies at local scales have demonstrated the importance of fruit during fall migratory stopover and (c) fruiting phenology in North America is occurring later over time while fall migration is advancing, the potential for mismatch between fall fruiting and bird migration phenology urgently needs further investigation., (© 2024 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Learned behavior, a fundamental adaptive trait in fluctuating environments, is shaped by species-specific constraints. This phenomenon is evident in songbirds, which acquire their species-specific songs through vocal learning. To explore the neurogenetic mechanisms underlying species-specific song learning, we generated F 1 hybrid songbirds by crossing Taeniopygia guttata with Aidemosyne modesta . These F 1 hybrids demonstrate expanded learning capacities, adeptly mimicking songs from both parental species and other heterospecific songs more extensively than their parental counterparts. Despite the conserved size of brain regions and neuron numbers in the neural circuits for song learning and production, single-cell transcriptomics reveals distinctive transcriptional characteristics in the F 1 hybrids, especially in vocal-motor projection neurons. These neurons exhibit enrichment for nonadditively expressed genes, particularly those related to ion channel activity and cell adhesion, which are associated with the degree of song learning among F 1 individuals. Our findings provide insights into the emergence of altered learning capabilities through hybridization, linked to cell type-specific transcriptional changes.

Fire suppression has negatively impacted thousands of acres of private and public lands in the United States. As a case study, the New Jersey Pine Barrens (NJPB) are a disturbance driven ecosystem that is experiencing serious ecological implications due to a loss of traditional forest thinning activities such as harvesting for forest products or thinning for wildfire fuel-load reduction measures coupled with a long-standing philosophy of fire suppression and dormant-season prescribed burning. Dense closed-canopy forest conditions, dissimilar to historic open-canopy forests of the NJPB, have reduced abundance and diversity of certain flora and fauna, including regionally imperiled breeding birds. In recent years, active forest stewardship (e.g., thinning, clear-cutting, and burning) has occurred on private and some public lands within the NJPB; however, the impact of such management on breeding birds is unclear due to a paucity of research on this subject within the NJPB. During 2012, 2013, 2016, and 2017, we conducted repeat-visit point counts (n = 1,800) for breeding songbirds across 75 control and 75 treatment sites within the NJPB to assess the influence of forest structure at three strata levels (groundcover, midstory profile, and canopy) on breeding bird communities. Specifically, we constructed a hierarchical community abundance model within a Bayesian framework for Bird Conservation Region (BCR) 30 priority upland birds (n = 12) within three species suites: Forested Upland, Scrub-Shrub (or Young Forest), and Grassland. At the community level, we found a negative relationship between bird abundance and live tree basal area. At the BCR 30 suite level, we found no relationship between Forested Upland suite-level abundance and any of the measured covariates; however, we found a negative relationship between percentage of woody groundcover and Scrub-Shrub suite-level abundance, and negative relationship between horizontal visual obstruction at 2 m above ground level and Grassland suite-level abundance. Furthermore, the two latter species suites exhibited a strong negative relationship with basal area. We recommend active forest stewardship that specifically targets opening the canopy to achieve basal areas between ~0-15 m2/ha via selective thinning, shelter cutting, and small-scale clear cutting. Mechanical treatment and prescribed burning would produce such conditions and have the added benefit of reducing fuel loads across this ~4,500 km2 landscape as well as assisting in carbon defense strategies for the region., Competing Interests: NO authors have competing interests, (Copyright: © 2024 Williams et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Population differences in socially learned mating signals like oscine birdsong are particularly vulnerable to breakdown through dispersal. 1 Despite this challenge, geographic variation in learned signals is ubiquitous. 2 A proposed explanation for this pattern is that birds express predispositions to selectively learn and produce population-typical songs. 3 , 4 , 5 While experimental studies on lab-reared birds have shown the existence of within-species learning predispositions, 6 , 7 , 8 , 9 , 10 it remains unclear whether and how learning predispositions influence song acquisition in the wild. Here, we investigated innate song learning predispositions in wild pied flycatchers (Ficedula hypoleuca) by measuring the songs of individuals translocated as eggs from a Dutch population to a breeding population in Sweden. We compared the songs of the adult males hatched from these translocated eggs with those from the ancestral and receiving populations. Songs of translocated males closely resemble the local Swedish songs to which they were exposed during development, supporting the importance of social learning. However, translocated males selectively learned those local Swedish song elements that sound the most "Dutch-like." As a result, their songs are significantly shifted toward those of the ancestral Dutch population. This suggests that innate learning predispositions track ongoing song evolution in wild populations of pied flycatchers. We propose that as songs continue to diverge over time, this coevolutionary relationship between song and learning predispositions may contribute to the emergence of incipient pre-mating barriers., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

Knowledge of the effect of harsh weather on the phenotypic traits of organisms is essential for understanding the environmental influence on phenotype evolution and holds implications for predicting how species respond to current climate change. For many birds, harsh weather in winter often imposes a strong selective effect on their survival, and only the individuals with certain phenotypes may survive. However, whether the selective effect on phenotype varies with winter weather conditions has been poorly investigated. Here, we explored the selective effect of winter weather on black-throated tit's (Aegithalos concinnus) morphological traits under winters with and without severe snowstorms. We found that for males, the sizes of their bills, heads and wings significantly affected their overwinter survival, but the effects varied with winter conditions. In relatively benign winters, males with smaller bill depths, smaller bill surface areas, and greater head lengths survived better; whereas, in winters with severe snowstorms, a reverse pattern was found. This phenomenon was likely driven by selection pressures from heat retention and foraging requirements, with their relative importance depending on winter conditions. Additionally, wing length was positively correlated with male survival and the relationship was stronger in harsher winters, which was probably due to longer wings' higher flight efficiency in adverse weather. By contrast, we found no correlation between morphological traits and survival in females. These results suggest a sex-specific and condition-dependent selective effect of environment on bird phenotypes, implying complicated interactions between different selection pressures and phenotype evolution., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Synchronous variation in demographic parameters across species increases the risk of simultaneous local extinction, which lowers the probability of subsequent recolonization. Synchrony therefore tends to destabilize meta-populations and meta-communities. Quantifying interspecific synchrony in demographic parameters, like abundance, survival, or reproduction, is thus a way to indirectly assess the stability of meta-populations and meta-communities. Moreover, it is particularly informative to identify environmental drivers of interspecific synchrony because those drivers are important across species. Using a Bayesian hierarchical multisite multispecies mark-recapture model, we investigated temporal interspecific synchrony in annual adult apparent survival for 16 common songbird species across France for the period 2001-2016. Annual adult survival was largely synchronous among species (73%, 95% credible interval [47%-94%] of the variation among years was common to all species), despite species differing in ecological niche and life history. This result was robust to different model formulations, uneven species sample sizes, and removing the long-term trend in survival. Synchrony was also shared across migratory strategies, which suggests that environmental forcing during the 4-month temperate breeding season has a large-scale, interspecific impact on songbird survival. However, the strong interspecific synchrony was not easily explained by a set of candidate weather variables we defined a priori. Spring weather variables explained only 1.4% [0.01%-5.5%] of synchrony, while the contribution of large-scale winter weather indices may have been stronger but uncertain, accounting for 12% [0.3%-37%] of synchrony. Future research could jointly model interspecific variation and covariation in breeding success, age-dependent survival, and age-dependent dispersal to understand when interspecific synchrony in abundance emerges and destabilizes meta-communities., (© 2024 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Social play has been described in many animals. However, much of this social behaviour among birds, particularly in adults, is still relatively unexplored in terms of the environmental, psychological, and social dynamics of play. This paper provides an overview of what we know about adult social play in birds and addresses areas in which subtleties and distinctions, such as in play initiation and social organisation and its relationship to expressions of play, are considered in detail. The paper considers emotional, social, innovative, and cognitive aspects of play, then the environmental conditions and affiliative bonds, suggesting a surprisingly complex framework of criteria awaiting further research. Adult social play has so far been studied in only a small number of avian species, exclusively in those with a particularly large brain relative to body size without necessarily addressing brain functions and lateralization. When lateralization of brain function is considered, it can further illuminate a possibly significant relevance of play behaviour to the evolution of cognition, to management of emotions, and the development of sociality., Competing Interests: Declaration of Competing Interest The author has no conflicts of interest to declare., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Learning and execution of complex motor skills are often modulated by sensory feedback and contextual cues arriving across multiple sensory modalities. Vocal motor behaviors, in particular, are primarily influenced by auditory inputs, both during learning and mature vocal production. The importance of auditory input in shaping vocal output has been investigated in several songbird species that acquire their adult song based on auditory exposure to a tutor during development. Recent studies have highlighted the influences of stimuli arriving through other sensory channels in juvenile song learning and in adult song production. Here, we review changes induced by diverse sensory stimuli during the song learning process and the production of adult song, considering the neuroethological significance of sensory channels in different species of songbirds. Additionally, we highlight advances, open questions, and possible future approaches for understanding the neural circuits that enable the multimodal shaping of singing behavior., Competing Interests: Declaration of competing interest All authors declare no competing interests., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Bird song is a classic example of a sexually selected trait, but much of the work relating individual song components to fitness has not accounted for song typically being composed of multiple, often-correlated components, necessitating a multivariate approach. We explored the role of sexual selection in shaping the complex male song of house wrens (Troglodytes aedon) by simultaneously relating its multiple components to fitness using multivariate selection analysis, which is widely used in insect and anuran studies but not in birds. The analysis revealed significant variation in the form and strength of selection acting on song across different selection episodes, from nest-site defense to recruitment of offspring to the breeding population. Males that sang more song typically employed in close communication sired more offspring that were subsequently recruited to the breeding population than those that sang more far-communication song. However, this relationship was not consistent across earlier selection episodes, as evidenced by non-linear selection acting on these song components in other contexts. Collectively, our results present a complex picture of multivariate selection on male song structure that would not be evident using univariate approaches and suggest possible trade-offs within and among song components at different points of the breeding season., (© The Author(s) 2024. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE). All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Global climate change involves various aspects of climate, including precipitation changes and declining surface wind speeds, but studies investigating biological responses have often focused on the impacts of rising temperatures. Additionally, related long-term studies on bird reproduction tend to concentrate on breeding onset, even though other aspects of breeding could also be sensitive to the diverse weather aspects. This study aimed to explore how multiple aspects of breeding (breeding onset, hatching delay, breeding season length, clutch size, fledgling number) were associated with different weather components. We used an almost four-decade-long dataset to investigate the various aspects of breeding parameters of a collared flycatcher (Ficedula albicollis) population in the Carpathian Basin. Analyses revealed some considerable associations, for example, breeding seasons lengthened with the amount of daily precipitation, and clutch size increased with the number of cool days. Parallel and opposing changes in the correlated pairs of breeding and weather parameters were also observed. The phenological mismatch between prey availability and breeding time slightly increased, and fledgling number strongly decreased with increasing mistiming. Our results highlighted the intricate interplay between climate change and the reproductive patterns of migratory birds, emphasizing the need for a holistic approach. The results also underscored the potential threats posed by climate change to bird populations and the importance of adaptive responses to changing environmental conditions., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Several metabolic hormones signal an organism's energy balance to the brain and modulate feeding behaviours accordingly. These metabolic signals may also regulate other behaviour related to energy balance, such as food caching or hoarding. Ghrelin is one such hormone, but it appears to exert different effects on appetite and fat levels in birds and mammals. Ghrelin treatment inhibits food intake and decreases fat stores in some bird species, but these effects may differ between acylated and unacylated (des-acyl) forms of ghrelin. The effect of ghrelin on food caching in birds has been examined in only one study, that found both leptin and unacylated ghrelin reduced food caching and mass gain in coal tits (Periparus ater). We expanded on this to test how both forms of ghrelin affect food caching and body composition in black-capped chickadees (Poecile atricapillus). We injected each bird with acylated ghrelin, unacylated ghrelin, and a saline control and then measured food caching every 20 min for two hours post-injection. We also measured body mass fat levels the day before, and after treatment using quantitative magnetic resonance (QMR). Contrary to prior work, we found no effects of either form of ghrelin on food caching, or body or fat mass. Future work is required to determine if the difference between our results and those of the prior study stems from species differences in response to ghrelin and/or in the motivation to cache food, or ghrelin effects being modulated by energy reserves., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

Despite extensive research on avian vocal learning, we still lack a general understanding of how and when this ability evolved in birds. As the closest living relatives of the earliest Passeriformes, the New Zealand wrens (Acanthisitti) hold a key phylogenetic position for furthering our understanding of the evolution of vocal learning because they share a common ancestor with two vocal learners: oscines and parrots. However, the vocal learning abilities of New Zealand wrens remain unexplored. Here, we test for the presence of prerequisite behaviors for vocal learning in one of the two extant species of New Zealand wrens, the rifleman (Acanthisitta chloris). We detect the presence of unique individual vocal signatures and show how these signatures are shaped by social proximity, as demonstrated by group vocal signatures and strong acoustic similarities among distantly related individuals in close social proximity. Further, we reveal that rifleman calls share similar phenotypic variance ratios to those previously reported in the learned vocalizations of the zebra finch, Taeniopygia guttata. Together these findings provide strong evidence that riflemen vocally converge, and though the mechanism still remains to be determined, they may also suggest that this vocal convergence is the result of rudimentary vocal learning abilities., (© 2024. The Author(s).)

Mixed-species groups and aggregations are quite common and may provide substantial fitness-related benefits to group members. Individuals may benefit from the overall size of the mixed-species group or from the diversity of species present, or both. Here we exposed mixed-species flocks of songbirds (Carolina chickadees, Poecile carolinensis, tufted titmice, Baeolophus bicolor, and the satellite species attracted to these two species) to three different novel feeder experiments to assess the influence of mixed-species flock size and composition on ability to solve the feeder tasks. We also assessed the potential role of habitat density and traffic noise on birds' ability to solve these tasks. We found that likelihood of solving a novel feeder task was associated with mixed-species flock size and composition, though the specific social factor involved depended on the particular species and on the novel feeder. We did not find an influence of habitat density or background traffic noise on likelihood of solving novel feeder tasks. Overall, our results reveal the importance of variation in mixed-species group size and diversity on foraging success in these songbirds., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Freeberg et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Substantial progress has been made in understanding the genetic architecture of phenotypes involved in a variety of evolutionary processes. Behavioral genetics remains, however, among the least understood. We explore the genetic architecture of spatial cognitive abilities in a wild passerine bird, the mountain chickadee (Poecile gambeli). Mountain chickadees cache thousands of seeds in the fall and require specialized spatial memory to recover these caches throughout the winter. We previously showed that variation in spatial cognition has a direct effect on fitness and has a genetic basis. It remains unknown which specific genes and developmental pathways are particularly important for shaping spatial cognition. To further dissect the genetic basis of spatial cognitive abilities, we combine experimental quantification of spatial cognition in wild chickadees with whole-genome sequencing of 162 individuals, a new chromosome-scale reference genome, and species-specific gene annotation. We have identified a set of genes and developmental pathways that play a key role in creating variation in spatial cognition and found that the mechanism shaping cognitive variation is consistent with selection against mildly deleterious non-coding mutations. Although some candidate genes were organized into connected gene networks, about half do not have shared regulation, highlighting that multiple independent developmental or physiological mechanisms contribute to variation in spatial cognitive abilities. A large proportion of the candidate genes we found are associated with synaptic plasticity, an intriguing result that leads to the hypothesis that certain genetic variants create antagonism between behavioral plasticity and long-term memory, each providing distinct benefits depending on ecological context., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Temporal changes in environmental conditions may play a major role in the year-to-year variation in fitness consequences of behaviours. Identifying environmental drivers of such variation is crucial to understand the evolutionary trajectories of behaviours in natural contexts. However, our understanding of how environmental variation influences behaviours in the wild remains limited. Using data collected over 14 breeding seasons from a collared flycatcher (Ficedula albicollis) population, we examined the effect of environmental variation on the relationship between survival and risk-taking behaviour, a highly variable behavioural trait with great evolutionary and ecological significance. Specifically, using annual recapture probability as a proxy of survival, we evaluated the specific effect of predation pressure, food availability, and mean temperature on the relationship between annual recapture probability and risk-taking behaviour (measured as flight initiation distance [FID]). We found a negative trend, as the relationship between annual recapture probability and FID decreased over the study years and changed from positive to negative. Specifically, in the early years of the study, risk-avoiding individuals exhibited a higher annual recapture probability, whereas in the later years, risk-avoiders had a lower annual recapture probability. However, we did not find evidence that any of the considered environmental factors mediated the variation in the relationship between survival and risk-taking behaviour., (© The Author(s) 2024. Published by Oxford University Press on behalf of the European Society of Evolutionary Biology. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

In this work, we used a dynamical system derived from an avian vocal production model to generate synthetic songs that mimic the Zonotrichia capensis songs. We confirmed that these synthetic renditions elicited behavioral responses similar to those evoked by real songs in wild songbirds of the same species. Specifically, we observed an increase in the singing rate of individual birds when a playback device was introduced into their territories. The success of our approach instills confidence in the hypotheses underpinning the model and provides a valuable tool for investigating a wide range of biological questions.

Migratory songbirds may navigate by extracting positional information from the geomagnetic field, potentially with a magnetic-particle-based receptor. Previous studies assessed this hypothesis experimentally by exposing birds to a strong but brief magnetic pulse aimed at remagnetizing the particles and evoking an altered behaviour. Critically, such studies were not ideally designed because they lacked an adequate sham treatment controlling for the induced electric field that is fundamentally associated with a magnetic pulse. Consequently, we designed a sham-controlled magnetic-pulse experiment, with sham and treatment pulse producing a similar induced electric field, while limiting the sham magnetic field to a value that is deemed insufficient to remagnetize particles. We tested this novel approach by pulsing more than 250 wild, migrating European robins ( Erithacus rubecula ) during two autumn seasons. After pulsing them, five traits of free-flight migratory behaviour were observed, but no effect of the pulse could be found. Notably, one of the traits, the migratory motivation of adults, was significantly affected in only one of the two study years. Considering the problem of reproducing experiments with wild animals, we recommend a multi-year approach encompassing large sample size, blinded design and built-in sham control to obtain future insights into the role of magnetic-particle-based magnetoreception in bird navigation.

Artificial light at night (ALAN) widely affects wildlife by blurring light-dark differences, including transitions such as sunrise and sunset, thereby affecting regulation of diel rhythms. As a result, activity onsets in many wild diurnal songbirds advance under ALAN. From chronobiological studies, it is known that the direction and strength of the response to light depends on when during the night exposure takes place. However, these experiments are mostly done under continuous light conditions, when animals have free-running rhythms. It remains unclear whether phase-dependence also holds in entrained, wild songbirds; i.e., does the effect of ALAN on activity patterns differ between exposure in the morning compared to the evening? This information is essential to assess the effects of mitigation measures by limiting ALAN to selected times of the night. We exposed incubating great tits (Parus major) inside the nest-box to 4 h of dim light, of which 1 h overlapped with dawn before sunrise or dusk after sunset. We found a small advancing effect of morning-light on activity onset and of evening-light on offset compared to dark controls but not vice versa. Breeding success and chick condition were unaffected by the light treatments. However, light-treated females had lower weights 9-18 days after the end of the treatment compared to the controls, independent of whether ALAN occurred in the morning or the evening, indicating possible costs of ALAN. Despite the weak behavioral response, ALAN might have affected the females' circadian clock or physiology resulting in lower body condition., (© 2024 The Authors. Journal of Experimental Zoology Part A: Ecological and Integrative Physiology published by Wiley Periodicals LLC.)

Migratory bird populations are declining globally at alarming rates. Non-breeding site conditions affect breeding populations, but generalising non-breeding habitat conditions over large spatial regions cannot address potential fine-scale differences across landscapes or local populations. Plumage characteristics can mediate the effects of environmental conditions on individual fitness. However, whether different phenotypes use distinctive non-breeding sites, and whether they respond to non-breeding site conditions differently remains largely unknown. Stable isotopes (δ 13 C, δ 15 N, δ 2 H) of inert tissues are useful to infer habitat characteristics and geographic origins where those tissues were grown. We collected winter-grown feathers from pied flycatchers (Ficedula hypoleuca) on their breeding grounds over several years from males whose dorsal plumage colouration ranged continuously from brown to black and assessed their stable isotope values as proxies of local habitat conditions. Based on feather δ 2 H profiles we found that browner males spent their non-breeding season in drier habitats than black males. Assignment to origin analysis shows potential regional non-breeding ground separation between differently coloured males. High within-individual repeatability of both δ 13 C and δ 15 N indicate the pied flycatcher males return yearly to similar areas. Blacker males were more likely to return to the breeding grounds after dry years compared with brown males. The opposite was found in wet years. Our study demonstrates that different phenotypes are exposed to different non-breeding site conditions which can differentially affect individual survivorship. This has important ramifications for population dynamics under predicted climate change scenarios where especially brown phenotype pied flycatcher males may be under a risk of decreasing., (© 2024. The Author(s).)

In a recent study, Shvedov and colleagues used live two-photon imaging in transgenic zebra finches to reveal migration patterns of neuroblasts through the complex environment of the postembryonic brain. This study highlights the value of ubiquitin C/green fluorescent protein (UBC-GFP) transgenic zebra finches in studying adult neurogenesis and advances our understanding of dispersed long-distance neuronal migration in the adult brain, shedding light on this understudied phenomenon., Competing Interests: Declaration of interests The author declares no competing interests in relation to this work., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Black-capped chickadees ( Poecile atricapillus ) and other species that feed at bird feeders balance the benefit of easy foraging with the added risk of predation. Individual birds respond differently to risky situations, and these differences have been attributed to the birds' personalities, which researchers commonly assess with an "open-field" behavioral assay. However, these behavioral assays in birds have not been compared to behavior in the wild in the context of foraging in the presence of a predator (i.e., risk-taking behavior). We color-banded chickadees in a wild population and conducted behavioral assays in the field. We later used foraging trials to investigate these color-banded individuals' responses to a predator (Cooper's hawk, Accipiter cooperii ) model or a series of Cooper's hawk calls. We found that foraging black-capped chickadees responded more strongly to the presence of a predator model than to predator calls. Individual birds differed in their responses, and the behavioral assays (activity and exploration) predicted individual behavior in the wild during the foraging experiments. Activity and exploration assay scores were only weakly related, suggesting these two assays represent different traits. Both highly active birds and fast explorers exhibited some reluctance to visit the feeder (either reduced number of visits or greater latency to visit) when the predator model was present, a relationship that was somewhat unexpected. Our results suggest that standard behavioral assays predict behavior in the wild, but care should be taken when generalizing among species and studies. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Birdsongs are among the most distinctive animal signals. Their evolution is thought to be shaped simultaneously by habitat structure and by the constraints of morphology. Habitat structure affects song transmission and detectability, thus influencing song (the acoustic adaptation hypothesis), while body size and beak size and shape necessarily constrain song characteristics (the morphological constraint hypothesis). Yet, support for the acoustic adaptation and morphological constraint hypotheses remains equivocal, and their simultaneous examination is infrequent. Using a phenotypically diverse Australasian bird clade, the honeyeaters (Aves: Meliphagidae), we compile a dataset consisting of song, environmental, and morphological variables for 163 species and jointly examine predictions of these two hypotheses. Overall, we find that body size constrains song frequency and pace in honeyeaters. Although habitat type and environmental temperature influence aspects of song, that influence is indirect, likely via effects of environmental variation on body size, with some evidence that elevation constrains the evolution of song peak frequency. Our results demonstrate that morphology has an overwhelming influence on birdsong, in support of the morphological constraint hypothesis, with the environment playing a secondary role generally via body size rather than habitat structure. These results suggest that changing body size (a consequence of both global effects such as climate change and local effects such as habitat transformation) will substantially influence the nature of birdsong.

Moult and migration are energetically demanding and require adequate nutrition. In some species, individuals may interrupt their fall migration to moult at discrete stopover locations outside of their breeding grounds (i.e., moult-migration) leading to competing nutritional demands for moult and migration. Here, we use DNA barcoding of fecal samples to compare the diet of moulting and actively migrating (post-moult) Swainson's Thrushes (Catharus ustulatus) and Tennessee Warblers (Leiothlypis peregrina) during their fall migration stopover at a large urban greenspace in Montreal, Canada. Diet differed according to moult status, species, and seasonality. Swainson's Thrushes had a broad diet with frequent detections of both insects and berry-producing shrubs; while detections in Tennessee Warblers' diets were mainly arthropods. For both species, more actively migrating individuals consumed fleshy-fruiting plants than moulting individuals. A higher proportion of moulting birds consumed arthropods compared to active migrants, due to either arthropod availability or a dietary preference for proteinaceous foods to grow feathers. Both species and moult classes consumed more native plants than non-native plants later in the season. We show the importance of managing urban greenspaces with native plants and diverse food sources that can provide for the different dietary needs of migratory birds., (© 2024. The Author(s).)

Cooperation may emerge from intrinsic factors such as social structure and extrinsic factors such as environmental conditions. Although these factors might reinforce or counteract each other, their interaction remains unexplored in animal populations. Studies on multilevel societies suggest a link between social structure, environmental conditions and individual investment in cooperative behaviours. These societies exhibit flexible social configurations, with stable groups that overlap and associate hierarchically. Structure can be seasonal, with upper-level units appearing only during specific seasons, and lower-level units persisting year-round. This offers an opportunity to investigate how cooperation relates to social structure and environmental conditions. Here, we study the seasonal multilevel society of superb fairy-wrens ( Malurus cyaneus ), observing individual responses to experimental playback of conspecific distress calls. Individuals engaged more in helping behaviour and less in aggressive/territorial song during the harsher non-breeding season compared to the breeding season. The increase in cooperation was greater for breeding group members than for members of the same community, the upper social unit, comprised of distinct breeding groups in association. Results suggest that the interaction between social structure and environmental conditions drives the seasonal switch in cooperation, supporting the hypothesis that multilevel societies can emerge to increase cooperation during harsh environmental conditions.

Seasonal migration is a widespread behavior relevant for adaptation and speciation, yet knowledge of its genetic basis is limited. We leveraged advances in tracking and sequencing technologies to bridge this gap in a well-characterized hybrid zone between songbirds that differ in migratory behavior. Migration requires the coordinated action of many traits, including orientation, timing, and wing morphology. We used genetic mapping to show these traits are highly heritable and genetically correlated, explaining how migration has evolved so rapidly in the past and suggesting future responses to climate change may be possible. Many of these traits mapped to the same genomic regions and small structural variants indicating the same, or tightly linked, genes underlie them. Analyses integrating transcriptomic data indicate cholinergic receptors could control multiple traits. Furthermore, analyses integrating genomic differentiation further suggested genes underlying migratory traits help maintain reproductive isolation in this hybrid zone., Competing Interests: Competing interests statement:The authors declare no competing interest.

Songs constitute a complex system of vocal signals for inter-individual communication in songbirds. Here, we elucidate the flexibility which songbirds exhibit in the organizing and sequencing of syllables within their songs. Utilizing a newly devised song decoder for quasi-real-time annotation, we execute an operant conditioning paradigm, with rewards contingent upon specific syllable syntax. Our analysis reveals that birds possess the capacity to modify the contents of their songs, adjust the repetition length of particular syllables and employing specific motifs. Notably, birds altered their syllable sequence in a goal-directed manner to obtain rewards. We demonstrate that such modulation occurs within a distinct song segment, with adjustments made within 10 minutes after cue presentation. Additionally, we identify the involvement of the parietal-basal ganglia pathway in orchestrating these flexible modulations of syllable sequences. Our findings unveil an unappreciated aspect of songbird communication, drawing parallels with human speech., (© 2024. The Author(s).)

Vocal rhythm plays a fundamental role in sexual selection and species recognition in birds, but little is known of its genetic basis due to the confounding effect of vocal learning in model systems. Uncovering its genetic basis could facilitate identifying genes potentially important in speciation. Here we investigate the genomic underpinnings of rhythm in vocal non-learning Pogoniulus tinkerbirds using 135 individual whole genomes distributed across a southern African hybrid zone. We find rhythm speed is associated with two genes that are also known to affect human speech, Neurexin-1 and Coenzyme Q8A. Models leveraging ancestry reveal these candidate loci also impact rhythmic stability, a trait linked with motor performance which is an indicator of quality. Character displacement in rhythmic stability suggests possible reinforcement against hybridization, supported by evidence of asymmetric assortative mating in the species producing faster, more stable rhythms. Because rhythm is omnipresent in animal communication, candidate genes identified here may shape vocal rhythm across birds and other vertebrates., (© 2024. The Author(s).)

Birds remodel their flight muscle metabolism prior to migration to meet the physiological demands of migratory flight, including increases in both oxidative capacity and defence against reactive oxygen species. The degree of plasticity mediated by changes in these mitochondrial properties is poorly understood but may be explained by two non-mutually exclusive hypotheses: variation in mitochondrial quantity or in individual mitochondrial function. We tested these hypotheses using yellow-rumped warblers (Setophaga coronata), a Nearctic songbird which biannually migrates 2000-5000 km. We predicted higher flight muscle mitochondrial abundance and substrate oxidative capacity, and decreased reactive oxygen species emission in migratory warblers captured during autumn migration compared with a short-day photoperiod-induced non-migratory phenotype. We assessed mitochondrial abundance via citrate synthase activity and assessed isolated mitochondrial function using high-resolution fluororespirometry. We found 60% higher tissue citrate synthase activity in the migratory phenotype, indicating higher mitochondrial abundance. We also found 70% higher State 3 respiration (expressed per unit citrate synthase) in mitochondria from migratory warblers when oxidizing palmitoylcarnitine, but similar H2O2 emission rates between phenotypes. By contrast, non-phosphorylating respiration was higher and H2O2 emission rates were lower in the migratory phenotype. However, flux through electron transport system complexes I-IV, II-IV and IV was similar between phenotypes. In support of our hypotheses, these data suggest that flight muscle mitochondrial abundance and function are seasonally remodelled in migratory songbirds to increase tissue oxidative capacity without increasing reactive oxygen species formation., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

The auditory sensitivity of a small songbird, the red-cheeked cordon bleu, was measured using the standard methods of animal psychophysics. Hearing in cordon bleus is similar to other small passerines with best hearing in the frequency region from 2 to 4 kHz and sensitivity declining at the rate of about 10 dB/octave below 2 kHz and about 35 dB/octave as frequency increases from 4 to 9 kHz. While critical ratios are similar to other songbirds, the long-term average power spectrum of cordon bleu song falls above the frequency of best hearing in this species., (© 2024 Acoustical Society of America.)

Trees release Herbivore-Induced Plant Volatiles (HIPVs) into the air in response to damage inflicted by insects. It is known that songbirds use those compounds to locate their prey, but more recently the idea emerged that songbirds could also use those odours as cues in their reproductive decisions, as early spring HIPVs may contain information about the seasonal timing and abundance of insects. We exposed pre-breeding great tits (Parus major) to the odours of caterpillar-infested trees under controlled conditions, and monitored reproduction (timing of egg laying, number of eggs, egg size) and two of its main hormonal drivers (testosterone and 17β-estradiol in males and females, respectively). We found that females exposed to HIPVs did not advance their laying dates, nor laid larger clutches, or larger eggs compared to control females. 17β-estradiol concentrations in females were also similar between experimental and control birds. However, males exposed to HIPVs had higher testosterone concentrations during the egg-laying period. Our study supports the hypothesis that insectivorous songbirds are able to detect minute amounts of plant odours. The sole manipulation of plant scents was not sufficient to lure females into a higher reproductive investment, but males increased their reproductive effort in response to a novel source of information for seasonal breeding birds., Competing Interests: Declaration of competing interest The authors have no competing interests to declare., (Copyright © 2024. Published by Elsevier Inc.)

A significant body of evidence indicates that climate change is influencing many aspects of avian ecology. Yet, how climate change is affecting, and is expected to influence some aspects of the breeding ecology of cavity-nesting birds remains uncertain. To explore the potential linkage between timing of first clutch, and the influence of ambient temperature on hatching success, we used Eastern Bluebird ( Sialia sialis ) nest records over a nine-year period from Alabama, USA. We investigated changes to annual clutch initiation dates, as well as variability in hatching success associated with ambient air temperatures during the incubation period. Using a simple linear model, we observed earlier annual egg laying dates over the nine years of this study with a difference of 24 days between earliest egg-laying date of the season. Daily temperature minima increased 2 °C across the nine-year time frame of this study. These data also indicate that Eastern Bluebird hatching success was the highest when mean ambient air temperature during incubation was between 19 °C and 24 °C (78%, as opposed to 69% and 68% above and below this temperature range, respectively). Our findings of increasing maxima, earlier maxima each year, and the lower minima of temperatures within our study area could expand the breadth of temperatures experienced by nesting Eastern Bluebirds possibly exposing them to temperatures outside of what promotes nesting success. These findings with a cavity-nesting bird highlight an optimal range of ambient temperatures associated with highest hatching success, conditions likely to be affected by climate change., Competing Interests: Declaration of conflicting interestsThe authors declared no potential conflicts of interest with respect to the research, authorship, and/or publication of this article.

Throughout the life of the adult songbird, neurons are recruited into brain regions important for song learning. Movies captured by Shvedov et al. demonstrate this dynamic process in the live animal, revealing the mechanisms of neuronal migration in the adult brain., Competing Interests: Declaration of interests The author declares no competing interests., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

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., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

Migration is an energetically challenging and risky life history stage for many animals, but could be supported by dietary choices en route, which may create opportunities to improve body and physiological condition. However, proposed benefits of diet shifts, such as between seasonally available invertebrates and fruits, have received limited investigation in free-living animals. We quantified diet composition and magnitude of autumn diet shifts over two time periods in two closely-related species of migratory songbirds on stopover in the northeastern U.S. (Swainson's thrush [Catharus ustulatus], long-distance migrant, N = 83; hermit thrush [C. guttatus], short-distance migrant, N = 79) and used piecewise structural equation models to evaluate the relationships among (1) migration timing, (2) dietary behavior, and (3) morphometric and physiological condition indices. Tissue isotope composition indicated that both species shifted towards greater fruit consumption. Larger shifts in recent weeks corresponded to higher body condition in Swainson's, but not hermit thrushes, and condition was more heavily influenced by capture date in Swainson's thrushes. Presence of "high-antioxidant" fruits in fecal samples was unrelated to condition in Swainson's thrushes and negatively related to multiple condition indices in hermit thrushes, possibly indicating the value of fruits during migration is related more to their energy and/or macronutrient content than antioxidant content. Our results suggest that increased frugivory during autumn migration can support condition, but those benefits might depend on migration strategy: a longer-distance, more capital-dependent migration strategy could require stricter regulation of body condition aided by increased fruit consumption., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Males usually come into conflict due to competition for territories and females. However, interference competition can also occur between males of congeneric species when their ecological requirements are overlapping. Using acoustic playback experiments, we investigated male-male interactions within and between Grasshopper (Locustella naevia; GW) and River Warbler (L. fluviatilis; RW). Our objective was to evaluate the song and behavioural response of tested males of both species to conspecific song stimuli in order to compare this with the response to congeneric stimulus, based on which we could assess whether these two commonly co-existing species show interspecific territorialism. A total of nine GW and 11 RW males were tested in May and June 2019 in western Slovakia. The ability to differentiate between the heterospecific (control), congeneric, and conspecific stimuli was similar between the two species. Conspecific playback elicited the strongest non-vocal response and a significant change in vocalization. The GW males shortened the songs, while the RW males shortened the songs and also increased their syllable rate. The congeneric playback elicited a lower intensity of behavioural response than conspecific playback and no change in vocalization in either species. We conclude that interspecific interference competition between GW and RW is rather low, suggesting that the species' ecological requirements are separated, although these two congeneric species commonly share habitat., Competing Interests: Declaration of Competing Interest None., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Neuron migration is a key phase of neurogenesis, critical for the assembly and function of neuronal circuits. In songbirds, this process continues throughout life, but how these newborn neurons disperse through the adult brain is unclear. We address this question using in vivo two-photon imaging in transgenic zebra finches that express GFP in young neurons and other cell types. In juvenile and adult birds, migratory cells are present at a high density, travel in all directions, and make frequent course changes. Notably, these dynamic migration patterns are well fit by a superdiffusive model. Simulations reveal that these superdiffusive dynamics are sufficient to disperse new neurons throughout the song nucleus HVC. These results suggest that superdiffusive migration may underlie the formation and maintenance of nuclear brain structures in the postnatal brain and indicate that transgenic songbirds are a useful resource for future studies into the mechanisms of adult neurogenesis., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

We investigated within- and between-individual song variation and song-based neighbour-stranger discrimination in a non-learning bird species, the blue-headed wood-dove (Turtur brehmeri), which inhabits lowland rainforests of West and Central Africa. We found that songs of this species are individually specific and have a high potential for use in individual recognition based on the time-frequency pattern of note distribution within song phrases. To test whether these differences affect behaviour, we conducted playback experiments with 19 territorial males. Each male was tested twice, once with the songs of a familiar neighbour and once with the songs of an unfamiliar stranger. We observed that males responded more aggressively to playback of a stranger's songs: they quickly approached close to the speaker and spent more time near it. However, no significant differences between treatments were observed in the vocal responses. In addition, we explored whether responses differed based on the song frequency of the focal male and/or that of the simulated intruder (i.e., playback), as this song parameter is inversely related to body size and could potentially affect males' decisions to respond to other birds. Song frequency parameters (of either the focal male or the simulated intruder) had no effect on the approaching response during playback. However, we found that the pattern of response after playback was significantly affected by the song frequency of the focal male: males with lower-frequency songs stayed closer to the simulated intruder for a longer period of time without singing, while males with higher-frequency songs returned more quickly to their initial song posts and resumed singing. Together, these results depict a consistently strong response to strangers during and after playback that is dependent on a male's self-assessment rather than assessment of a rival's strength based on his song frequency. This work provides the first experimental evidence that doves (Columbidae) can use songs for neighbour-stranger discrimination and respond according to a "dear enemy" scheme that keeps the cost of territory defence at a reasonable level., (© 2024. The Author(s).)

Migratory birds possess remarkable accuracy in orientation and navigation, which involves various compass systems including the magnetic compass. Identifying the primary magnetosensor remains a fundamental open question. Cryptochromes (Cry) have been shown to be magnetically sensitive, and Cry4a from a migratory songbird seems to show enhanced magnetic sensitivity in vitro compared to Cry4a from resident species. We investigate Cry and their potential involvement in magnetoreception in a phylogenetic framework, integrating molecular evolutionary analyses with protein dynamics modelling. Our analysis is based on 363 bird genomes and identifies different selection regimes in passerines. We show that Cry4a is characterized by strong positive selection and high variability, typical characteristics of sensor proteins. We identify key sites that are likely to have facilitated the evolution of an optimized sensory protein for night-time orientation in songbirds. Additionally, we show that Cry4 was lost in hummingbirds, parrots and Tyranni (Suboscines), and thus identified a gene deletion, which might facilitate testing the function of Cry4a in birds. In contrast, the other avian Cry (Cry1 and Cry2) were highly conserved across all species, indicating basal, non-sensory functions. Our results support a specialization or functional differentiation of Cry4 in songbirds which could be magnetosensation.