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Birdsong is a longstanding model system for studying evolution, and has recently emerged as a measure of biodiversity loss due to deforestation and climate change. Here, we collected and analyzed high quality song recordings from seven species in the family Estrildidae. We measured the acoustic features of syllables and then used dimensionality reduction and machine learning classifiers to identify features that accurately assigned syllables to species. Species differences were captured by the first 3 principal components, corresponding to basic spectral features, spectral shape, and spectrotemporal features. We then identified the measured features underlying classification accuracy. We found that fundamental frequency, mean frequency, spectral flatness, and syllable duration were the most informative features for species identification. Next, we tested whether specific acoustic features of species’ songs predicted phylogenetic distance. We found significant phylogenetic signal in syllable spectral features, but not in spectral shape or spectrotemporal features. Results indicate that spectral features are more constrained by species’ genetics than are other features, and are the best signal features for identifying species from song recordings. The absence of phylogenetic signal in spectral shape and spectrotemporal features suggests that these song features are labile, reflecting learning-processes and individual recognition.

Eggs are critically important for avian reproduction as all birds are oviparous. Accordingly, the recognition and care of own eggs represent the cornerstones of avian breeding, whereas the elimination of foreign objects, including brood-parasitic eggs and non-egg items from the nest are known to also increase fitness by refocusing incubation effort on the parents' own eggs. But egg recognition also plays a role in some avian obligate brood parasites' reproductive strategy through the pecking of already present eggs in the hosts' clutch to reduce nestmate competition with the parasite's own hatchling. Here we tested egg shape recognition in this parasitic egg-pecking context by exposing two different series of 3D printed models to captive obligate brood-parasitic shiny cowbirds ( Molothrus bonariensis ) in artificial nests. Natural egg-shaped models were pecked more often compared with increasingly thinner models, but there was no effect of increasing angularity on pecking rates, implying that a natural, rather than an artificial, range of variability elicited adaptive responses from parasitic cowbirds.

Classic models of vigilance assume instantaneous and sequential randomness in the scanning process, implying negative exponential distribution of interscan durations and no interdependence among successive interscans. We examined whether vigilance pattern by preening black-headed gulls, Chroicocephalus ridibundus, meets these assumptions. Out of 54 behavioural sequences, 50 departed from the expected negative exponential distribution, whereas the focal interscan duration was significantly affected by the interaction of the preceding scan and the interscan interval. These results reveal departures from randomness in the scanning process by gulls, which may be a consequence of the hunting strategies of their predators or due to the trade-off between the needs for feather maintenance and antipredator vigilance.

Avian eggshell pigmentation may provide information about a female’s physiological condition, in particular her state of oxidative balance. Previously we found that female House Wrens (Troglodytes aedon Vieillot, 1809) with lighter, less-maculated, and redder ground-colored shells were older and produced heavier offspring than females laying more-maculated and browner eggs. The strong pro-oxidant protoporphyrin is responsible for this species’ eggshell pigmentation, so differences in pigmentary coloration may be related to eggshell protoporphyrin content and reflect female oxidative balance and condition during egg formation. Therefore, we tested the assumption that egg-surface coloration is related to the amount of protoporphyrin in the shell matrix. We analyzed digital photographs of eggs to determine maculation coverage as a measure of the overall ground coloration of the egg and its red-, green-, and blue-channel pixel values. Pigments were then extracted from these same eggs and analyzed using high-performance liquid chromatography. There was a strong, positive relationship between eggshell redness and protoporphyrin content of eggshells, but no relationship between percent maculation and protoporphyrin content. Thus, when older, larger females deposit more protoporphyrin in their eggshells, this may reflect a tolerance for high levels of circulating protoporphyrin or an effective mechanism for off-loading protoporphyrin into the eggshell matrix.

Hybridization is a known source of morphological, functional and communicative signal novelty in many organisms. Although diverse mechanisms of established novel ornamentation have been identified in natural populations, we lack an understanding of hybridization effects across levels of biological scales and upon phylogenies. Hummingbirds display diverse structural colours resulting from coherent light scattering by feather nanostructures. Given the complex relationship between feather nanostructures and the colours they produce, intermediate coloration does not necessarily imply intermediate nanostructures. Here, we characterize nanostructural, ecological and genetic inputs in a distinctive Heliodoxa hummingbird from the foothills of eastern Peru. Genetically, this individual is closely allied with Heliodoxa branickii and Heliodoxa gularis , but it is not identical to either when nuclear data are assessed. Elevated interspecific heterozygosity further suggests it is a hybrid backcross to H. branickii . Electron microscopy and spectrophotometry of this unique individual reveal key nanostructural differences underlying its distinct gorget colour, confirmed by optical modelling. Phylogenetic comparative analysis suggests that the observed gorget coloration divergence from both parentals to this individual would take 6.6–10 My to evolve at the current rate within a single hummingbird lineage. These results emphasize the mosaic nature of hybridization and suggest that hybridization may contribute to the structural colour diversity found across hummingbirds.

Nest-sharer avian brood parasites do not evict or otherwise kill host chicks, but instead inflict a range of negative effects on their nestmates that are mediated by interactions between the parasite and host life history traits. Although many of the negative fitness effects of avian brood parasitism are well documented across diverse host species, there remains a paucity of studies that have examined the impacts of parasitism across the entirety of host ontogeny (i.e., from when an egg is laid until independence). More specifically, few studies have examined the impact of brood parasitism on the pre- and post-fledging development, physiology, behavior, and survival of host offspring. To help fill this knowledge gap, we assessed the effects of brood parasitism by Brown-headed Cowbirds (Molothrus ater) across the ontogeny (incubation, nestling, and post-fledging period) of nine sympatrically breeding host species in central Illinois, USA; due to sample sizes, impacts on the post-fledging period were only examined in two of the nine species. Specifically, we examined the impact of brood parasitism on ontogenetic markers including the embryonic heart rate, hatching rate, nestling period length, nest survival, and offspring growth and development. Additionally, in species in which we found negative impacts of cowbird parasitism on host nestmate ontogeny, we examined whether the difference in adult size between parasites and their hosts and their hatching asynchrony positively predicted variation in host costs across these focal taxa. We found that costs of cowbird parasitism were most severe during early nesting stages (reduction in the host clutch or brood size) and were predicted negatively by host size and positively by incubation length. In contrast, we only found limited costs of cowbird parasitism on other stages of host ontogeny; critically, post-fledging survival did not differ between host offspring that fledged alongside cowbirds and those that did not. Our findings (i) highlight the direct costs of cowbird parasitism on host fitness, (ii) provide evidence for when (the stage) those costs are manifested, and (iii) may help to explain why many anti-cowbird defenses of hosts have evolved for protection from parasitism during the laying and incubation stages.

One of the most effective defenses of avian hosts against obligate brood parasites is the ejection of parasitic eggs from the nests. Despite the clear fitness benefits of this behavior, individuals within so-called “egg rejecter” host species still show substantial variation in their propensity to eliminate foreign eggs from the nest. We argue that this variation can be further understood by studying the physiological mechanisms of host responses to brood parasitic egg stimuli: independent lines of research increasingly support the hypothesis that stress-related physiological response to parasitic eggs may trigger egg rejection. The “stress-mediated egg rejection” hypothesis requires that hosts activate the stress-response when responding to parasitic eggs. We tested this prediction by experimentally parasitizing incubating American robins Turdus migratorius, an egg rejecter host to obligate brood parasitic brown-headed cowbirds Molothrus ater, with mimetic or non-mimetic model eggs. To assess the stress response, we measured the heart rate in incubating females immediately after experimental parasitism. We also measured plasma corticosterone and, in a subset of birds, used RNA-sequencing to analyze the expression of proopiomelanocortin (POMC), a precursor of adrenocorticotropic hormone (ACTH), two hours after experimental parasitism. We found that egg type had no effect on heart rate. Two hours following experimental parasitism, plasma corticosterone did not differ between the differently colored model egg treatments or between rejecter and accepter females within the non-mimetic treatment. However, females exposed to non-mimetic eggs showed an upregulation of POMC gene expression in the pituitary compared to females treated with mimetic eggs. Our findings suggest that parasitic eggs may activate the stress-related hypothalamic-pituitary-adrenal axis in an egg-rejecter host species, although the dynamics of this response are not yet understood.

For many birds, nest construction is a costly aspect of parental care, trading finite energetic resources between parental care and self-maintenance. For multi-brooded organisms with short breeding seasons, such as migratory passerines, repeated nest construction could be especially costly if the activity delays the onset of breeding attempts. Earlier studies on passerines that reuse nests between breeding seasons suggested that time lost to initial nest construction reduces seasonal reproductive output. However, costs associated with building new nests between breeding attempts, within the same breeding season, have largely been ignored. Here, we experimentally removed first nests, after fledging or failing, of Eastern Phoebes ( Sayornis phoebe), to evaluate how the annual onset of breeding and nest construction between breeding attempts affected parental investment into second attempts. We found that first egg laying date negatively predicted the probability of second breeding attempts, but experimental treatment (first nest removal vs. control) did not. Neither first egg laying date nor treatment statistically influenced any of the reproductive traits in second breeding attempts (clutch size, nestling body condition, and nestling growth rate). We conclude that in this species, second breeding attempts are limited by the initial onset of seasonal reproduction, and not by time lost to nest construction between breeding attempts.

The Prothonotary Warbler (Protonotaria citrea) is a nearctic-neotropical migratory songbird that breeds in forested swamps and riparian areas in the eastern-central United States and southern Ontario. It is the sole eastern North American wood-warbler that nests in cavities, the only species in the genus Protonotaria, and is one of the few hole-nesting hosts of the Brown-headed Cowbird (Molothrus ater), an obligate brood parasite. We present the whole genome sequence of this wood-warbler species. Illumina sequencing was performed on a genetic sample from a single female individual. The reads were assembled using a de novo method followed by a finishing step. The raw and assembled data is publicly available via Genbank: Sequence Read Archive (SRR19579445) and Assembly (JAOYNP000000000).

Obligate insect social parasites evolve traits to effectively locate and then exploit their hosts, whereas hosts have complex social behavioral repertoires, which include sensory recognition to reject potential conspecific intruders and heterospecific parasites. While social parasites and host behaviors have been studied extensively, less is known about how their sensory systems function to meet their specific selective pressures. Here, we compare investment in visual and olfactory brain regions in the paper wasp Polistes dominula, and its obligate social parasite P. sulcifer, to explore the links among sensory systems,brain and behavior. Our results show significant relative volumetric differences between these two closely related species, consistent with their very different life histories. Social parasites show proportionally larger optic lobes and central complex to likely navigate long-distance migrations and unfamiliar landscapes to locate the specific species of hosts they usurp. Contrastingly, hosts have larger antennal lobes and calyces of the mushroom bodies compared with social parasites, as predicted by their sensory means to maintain social cohesion via olfactory signals, allocate colony tasks, forage, and recognize conspecific and heterospecific intruders. Our work suggests how this tradeoff between visual and olfactory brain regions may facilitate different sensory adaptations needed to perform social and foraging tasks by the host, including recognition of parasites, or to fly long distances and successful host localizing by the social parasite.

The causes and consequences of interspecific variation in sex-specific contributions to animal parental care are relatively well understood during pregnancy or incubation and during offspring provisioning, but comparative patterns of sex-biased investment during nest-, den-, or other shelter-building have been almost completely overlooked. This is surprising because birthing shelters’ protective properties have important fitness consequences for both parents and offspring. Here, we address this gap in our knowledge by testing predictions concerning sex-specific contributions to avian nest building in more than 500 species of Western Palearctic birds in relation to the time available to breed and sex-specific reproductive effort, while also examining correlates with nesting site and nest structure. Using multivariate phylogenetic comparative and path analysis approaches, we found that, opposite to what had been predicted, species in which females build nests alone have shorter breeding seasons and breed at higher latitudes. In addition, species in which females lay larger clutch sizes and incubate eggs alone are more likely to have nests built by females alone, again countering predictions that reproductive contributions are not traded-off between the sexes. Finally, however, sex-specific nest building contributions were predictably related to nest site and structure, as species in which females built nests alone were more likely to have open cup nests relative to enclosed, domed nests of species in which both parents build. Our study provides important new insights, and generates several new questions for experimental research into the adaptive dynamics of sex-specific contributions prior or at the onset of parental care.

Generalist brood parasites that share nests with host nestlings can optimize resource acquisition from host parents by balancing the benefits that host nest-mates provide, including attracting increased provisions to the nest, against the costs of competing with the same host young over foster parental resources. However, it is unclear how parasitic chicks cope when faced with more nest-mates than are optimal for their survival upon hatching. We suggest that, in the obligate brood parasitic brown-headed cowbird ( Molothrus ater ), chicks use a niche construction strategy and reduce larger, more competitive host broods to maximize the parasites' survival to fledging. We experimentally altered brood sizes to test for Goldilocks principle patterns (i.e. a ‘just right’ intermediate brood size) of cowbird survival in nests of prothonotary warbler ( Protonotaria citrea ) hosts. We found that intermediate brood sizes of two host nestlings maximized cowbird fledging success relative to 0 or 4 host nest-mates at hatching. Specifically, cowbird nestlings lowered host brood sizes towards this optimum when placed in broods with more host nestlings. The results suggest that cowbirds reduce, but do not eliminate, host broods as a niche construction mechanism to improve their own probability of survival.

Avian obligate brood parasites rely on other species to raise their offspring. In turn, many brood parasite hosts have evolved defensive behaviors to reduce the costs of brood parasitism, yet the proximate bases underlying these defenses remain poorly understood. Recent studies regarding the potential endocrine mechanisms of foreign-egg rejection have implicated corticosterone as a physiological mediator of anti-parasitic defenses. For example, corticosterone is elevated in response to non-mimetic eggs in an egg rejecter thrush, the Eurasian blackbird (Turdus merula) and this hormone's suppression reduces egg rejection rates in the congeneric American robin (T. migratorius). American robins are also among the few host species of obligate brood parasitic brown-headed cowbirds (Molothrus ater) that readily reject foreign eggs from their nests. We non-invasively elevated corticosterone levels in incubating female robins by dissolving it in DMSO gel which was then applied onto eggs already in the clutch. Relative to controls treated with pure DMSO gel, corticosterone-treated female robins were more likely to reject a non-mimetic, cowbird-sized foreign egg (72 %) than control females (50 %) when accounting for the known effect of lower clutch sizes on greater egg rejection. Future studies are needed to assess the sensory and cognitive impact(s) of corticosterone, as well as other hormones essential for parental care, in this and other hosts' defense behaviors against avian brood parasitism.

In avian brood parasitism, parasites lay their eggs in the nests of hosts, and many hosts in the wild respond by eliminating or abandoning foreign eggs in their clutch. However, a limitation upon the study of proximate, especially physiological and experience-dependent cognitive mechanisms of egg rejection, has been the lack of a suitable model system in captivity. Here, we tested whether laboratory-kept ringneck doves

Obligate avian brood parasites depend on hosts for parental care, which in turn suffer fitness losses as a result of parasitism. Mechanisms by which brood parasitic cowbirds (Molothrus spp.) reduce host breeding success include the puncture (M. rufoaxillaris and M. bonariensis) or removal (M. ater) of the eggs of the host. Our working hypothesis is that the host eggs’ mechanical strength and their size and shape in species with higher frequency of parasitism covaries with the cowbird’s strategy to reduce host clutch size. Our results, obtained through phylogenetic analyses based on egg 2D geometric morphometry and eggshell mechanical and ultrastructural measurements, suggest that egg-puncturer behaviour has led to an increase in the strength of the host’s eggshell, which might make them more difficult to be pierced. We also characterized larger, more rounded and asymmetrical eggs in frequent hosts of M. ater, which might be more difficult to be removed. These interspecific host egg and shell traits were also positively affected by the frequency of parasitism, indicating that species-specific patterns of parasitic costs select for respective anti-parasitic defences in hosts.

This research investigates the extent and causal mechanisms of genetic population divergence in a poorly flighted passerine, the North Island Rifleman or Titipounamu (Acanthisitta chloris granti). While this species has a historically widespread distribution, anthropogenic forest clearance has resulted in a highly fragmented current distribution. We conducted analyses of mitochondrial DNA (COI and Control Region) and 12 nuclear DNA microsatellites to test for population divergence and estimate times of divergence. diyabc and biogeobears were then used to assess likely past dispersal scenarios based on both mtDNA and nDNA. The results reveal several significantly divergent lineages across the North Island of New Zealand and indicate that some populations have been isolated for extensive periods of time (0.7–4.9 mya). Modeling indicated a dynamic history of population connectivity, with a drastic restriction in gene flow between three geographic regions, followed by a more recent re‐establishment of connectivity. Our analyses indicate the dynamic influence of key geological and climatological events on the distribution of genetic diversity in this species, including support for the genetic impact of old biogeographic boundaries such as the Taupo Line and Cockayne's Line, rather than recent anthropogenic habitat fragmentation. These findings present a rare example of an avian species with a genetic history more like that of flightless taxa and so provide new general insights into vicariant processes affecting populations of passerines with limited dispersal., This research uncovers significant genetic divergence between populations of a basal passerine with restricted dispersal potential, New Zealand's North Island Rifleman. These results demonstrate the impact of ancient biogeographic events on the distribution of this species and present a rare example of an avian species with a genetic history more like that of flightless taxa. This work provides new insights into vicariant processes affecting populations of passerines with limited dispersal more broadly.

Many avian species are negatively impacted by obligate avian brood parasites, which lay their eggs in the nests of host species. The yellow warbler (Setophaga petechia), which is host to the brood-parasitic brown-headed cowbird (Molothrus ater), represents one of the best-replicated study systems assessing antiparasitic host defenses. Over 15 prior studies on yellow warblers have used model-presentation experiments, whereby breeding hosts are exposed to models of brown-headed cowbirds or other nest threats, to test for anti-parasitic defenses unique to this species. Here we present results from our own quasi-replication study of the yellow warbler/brown-headed cowbird system, which used a novel design compared to previous experiments by pivoting to conduct acoustic playback treatments only, rather than presenting visual models with or without calls. We exposed active yellow warbler nests to playbacks of brown-headed cowbird chatters (brood parasite), blue jay (Cyanocitta cristata; nest predator) calls, conspecific “seet” calls (a referential alarm call for brood parasitism risk), conspecific “chip” calls (a generic alarm call), or control wood thrush (Hylocichla mustelina; harmless heterospecific) songs during the incubation stage. Similar to previous studies, we found that female yellow warblers seet called more frequently in response to playbacks of both brood parasitic chatter calls and conspecific seet calls whereas they produced more chip calls in response to the playback of nest predator calls. In contrast, female yellow warblers approached all playbacks to similar distances, which was different from the proximity patterns seen in previous studies. Our study demonstrates the importance of both replicating, and also pivoting, experimental studies on nest defense behaviors, as differences in experimental design can elicit novel behavioral response patterns in the same species.

Avian obligate brood parasites lay their eggs in the nests of other species that may provide care for the foreign offspring. Brood parasitism often imparts substantial fitness losses upon host nestlings when they are raised alongside the typically more competitive, larger, and older parasitic chick(s). Whereas fitness costs due to reduced host offspring survival in parasitized broods have been studied in detail, the physiological changes in host nestlings caused by parasitic nestmate(s) are less well known. We compared prothonotary warbler (Protonotaria citrea) nestlings, a host of the nest-sharing brown-headed cowbird (Molothrus ater), in experimentally parasitized vs. non-parasitized broods. Our aim was to determine whether cohabitation with brood parasitic young impacted host nestling baseline corticosterone plasma concentrations, immune responses, body condition, and mortality. Corticosterone levels and body condition of host nestlings were similar between nests with or without a cowbird nestmate, whereas host immune responses were lower and nestling mortality was greater in parasitized broods, irrespective of variation in brood size or total brood mass. We detected no trade-offs of baseline corticosterone levels with either immune responses or with body condition. These results suggest that this host species' nestlings experience some adverse fitness-relevant physiological effects in parasitized broods, but are also resilient in other aspects when coping with brood parasitism.

In dense breeding colonies, and despite having no nest structure, common murres (or guillemots: Uria aalge) are still able to identify their own eggs. Each female murre's egg is thought to be recognized individually by the shell's avian‐perceivable traits. This is because the eggshells’ visible traits conform to expectations of the identity‐signaling hypothesis in that they show both high intraindividual repeatability and high interindividual variability. Identity signaling also predicts a lack of correlation between each of the putative multicomponent recognition traits, yielding no significant relationships between those eggshell traits that are generated by mutually exclusive physiological factors. Using a multivariate analysis across eggshell size and shape, avian‐perceivable background coloration, spot (maculation) shape, and spot density, we detected no unexpected statistical correlations between Icelandic common murre egg traits lacking known physiological or mathematical relationships with one another. These results biologically replicate the conclusions of a recent eggshell trait study of Canadian common murres using similar methodology. We also demonstrate the use of static correlations to infer identity signaling function without direct behavioral observations, which in turn may also be applied to rare or extinct species and provide valuable insight into otherwise unknown communicative and behavioral functions., A multivariate analysis of eggshells taken from an Icelandic population of common murre (or guillemot: Uria aalge) reveals that no unexpected correlations exist between avian‐perceivable eggshell traits. These results biologically replicate the conclusions of a recent eggshell trait study of Canadian common murres using the same methodology, as well as provide further evidence for the functionality of avian‐perceivable eggshell traits as identity signals in the common murre.

Generalist obligate brood parasites are excellent models for studies of developmental plasticity, as they experience a range of social and environmental variation when raised by one of their many hosts. Parasitic Brown-headed Cowbirds (Molothrus ater (Boddaert, 1783)) exhibit host-specific growth rates, yet Cowbird growth rates are not predicted by hosts’ incubation or brooding periods. We tested the novel “growth-tuning” hypothesis which predicts that total asynchrony between Cowbirds’ and hosts’ nesting periods results in faster parasitic growth in nests where host young fledge earlier than Cowbirds. We tested this prediction using previously published and newly added nestling mass data across diverse host species. Total nesting period asynchrony (summed across incubation and brooding stages) predicted Cowbird growth; 8-day-old Cowbirds were heavier in host nests with relatively shorter nesting periods. We further explored the drivers of variation in growth using mass measurements of Cowbirds in Song Sparrow (Melospiza melodia (A. Wilson, 1810)) and Red-winged Blackbird (Agelaius phoeniceus (Linnaeus, 1766)) nests. Our top models included host species (Cowbirds grew faster in Sparrow nests), numbers of nestmates (slowest when raised alone), and sex (males grew faster). These results confirm that multiple social and environmental factors predict directional patterns of developmental plasticity in avian generalist brood parasites.

Invasive species present an opportunity to test the association between selective forces and adaptive morphological traits because these species can experience rapid changes when introduced to new environments. One such invader is the common myna (Acridotheres tristis), a broadly ranging avian species that has been introduced on most continents and many oceanic islands. Here, we studied morphological variability in native and introduced populations of common mynas to identify the key environmental variables correlated with local phenotypic variation during biological invasions. In particular, we aimed to determine whether similar selective pressures acted on populations in the native range and the introduced ranges, and whether the same factors shaped morphological variation in different introduced populations. We recorded eight morphological measurements from 1,331 individuals across sites throughout most of the common myna’s range, as well as climatic, topographic and anthropogenic environmental data, sex and age of individuals, and years since local introduction. We found inconsistent signals of morphological adaptation in the native population, representing different patterns between females and males. Conversely, urbanization was a significant explanatory factor of phenotypic shifts in introduced populations, especially with regards to foraging-related traits. Moreover, we found that adaptation occurred differently across two focal introduced populations (Sydney, Australia and Israel). Therefore, caution is required in studies relating to evolutionary adaptations and predictions made regarding characteristics of invasive populations within a limited range of their introduced ranges. Our results also indicate a major role for urbanization in impacting the shape and size of foraging-related features in this broadly-distributed invasive species, emphasizing the impact of human-induced environmental changes upon the trajectories of biological invasions and, ultimately, their effects on native biodiversity.

When individuals groom or preen one another (allopreening), vulnerability and physical access allow for information about hierarchy and bonds to be exchanged, in addition to basic health benefits. During these interactions, individuals may also self-direct grooming/preening (autopreening). We investigate autopreening in a cooperatively breeding passerine bird, the Arabian babbler (Turdoides squamiceps) in the specific context of allopreening interactions. Auto- and allogrooming (the mammalian analogue behaviours) may occur in sequence together, with some species socially facilitating allogrooming by autogrooming. We here consider the possibility of similar associations between autopreening and allopreening. Using a two-year database of 35 individuals in 8 social groups, we assessed the social factors that predict the occurrence of autopreening and its timing in the context of allopreening dyad formation. We ask if there is a social dimension to this behaviour, and specifically find evidence that it may represent displacement and/or subordinate behaviour. The relative age of individuals in preening dyads and the behaviour of recipients prior to social approach significantly predicted the behaviour of recipients as the preening dyad was formed: recipients were less likely to autopreen before approaching older actors to begin an allopreening bout. When recipients autopreened, actors were more likely to make the approach. Recipients that did not autopreen or otherwise conspicuously display were significantly more likely to approach the actor without first autopreening. We suggest that autopreening in these bouts is a socially modulated behaviour representative of uncertainty and/or subordination.

Defending offspring incurs temporal and energetic costs and can be dangerous for the parents. Accordingly, the intensity of this costly behavior should reflect the perceived risk to the reproductive output. When facing costly brood parasitism by brown‐headed cowbirds (Molothrus ater), where cowbirds lay eggs in heterospecific nests and cause the hosts to care for their young, yellow warblers (Setophaga petechia) use referential “seet” calls to warn their mates of the parasitic danger. Yellow warblers of both sexes produce this call only in response to cowbirds or seet‐calling conspecifics. Seet calls are mainly produced during the laying and incubation stages of breeding, when risk of brood parasitism is highest, rather than during the nestling stage. On the other hand, general alarm calls (chips) are produced throughout the nesting cycle and are also used in conspecific interactions unrelated to nesting. We hypothesized that context shapes responses prior to breeding as well, such that yellow warblers without a mate and active nest would be less likely to respond to playbacks that simulate brood parasitism risk. To test this hypothesis, we presented playbacks of two nest threats, cowbirds (brood parasite) and blue jays (Cyanocitta cristata; nest predator), on territories of unmated male warblers (unpaired) and male warblers with a known mate (paired). We found that unpaired males were unresponsive toward playbacks indicating nest threats, whereas paired males were significantly more aggressive and vocal toward these playbacks compared to control playbacks. However, both paired and unpaired males were vocally responsive toward chip calls, which are informative for males regardless of pairing status. Male yellow warblers appear to adjust their responses during the earliest stages of breeding depending on the contextual relevance of specific threat stimuli, and together with prior studies, our work further supports that referential seet calls are associated with stage‐specific risk of brood parasitism.

Ongoing global change has had biologically impactful effects on the breeding phenology of both resident and migratory bird species, including avian hosts and their obligate brood parasites. We anal...

Understanding when learning begins is critical for identifying the factors that shape both the developmental course and the function of information acquisition. Until recently, sufficient development of the neural substrates for any sort of vocal learning to begin in songbirds was thought to be reached well after hatching. New research shows that embryonic gene activation and the outcome of vocal learning can be modulated by sound exposure in ovo . We tested whether avian embryos across lineages differ in their auditory response strength and sound learning in ovo , which we studied in vocal learning (Maluridae, Geospizidae) and vocal non-learning (Phasianidae, Spheniscidae) taxa. While measuring heart rate in ovo , we exposed embryos to (i) conspecific or heterospecific vocalizations, to determine their response strength, and (ii) conspecific vocalizations repeatedly, to quantify cardiac habituation, a form of non-associative learning. Response strength towards conspecific vocalizations was greater in two species with vocal production learning compared to two species without. Response patterns consistent with non-associative auditory learning occurred in all species. Our results demonstrate a capacity to perceive and learn to recognize sounds in ovo , as evidenced by habituation, even in species that were previously assumed to have little, if any, vocal production learning. This article is part of the theme issue ‘Vocal learning in animals and humans’.

The relationships between avian brood parasites and their hosts are widely recognised as model systems for studying coevolution. However, while most brood parasites are known to parasitise multiple species of host and hosts are often subject to parasitism by multiple brood parasite species, the examination of multispecies interactions remains rare. Here, we compile data on all known brood parasite–host relationships and find that complex brood parasite–host systems, where multiple species of brood parasites and hosts coexist and interact, are globally commonplace. By examining patterns of past research, we outline the disparity between patterns of network complexity and past research emphases and discuss factors that may be associated with these patterns. Drawing on insights gained from other systems that have embraced a multispecies framework, we highlight the potential benefits of considering brood parasite–host interactions as ecological networks and brood parasitism as a model system for studying multispecies interactions. Overall, our results provide new insights into the diversity of these relationships, highlight the stark mismatch between past research efforts and global patterns of network complexity, and draw attention to the opportunities that more complex arrangements offer for examining how species interactions shape global patterns of biodiversity.