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Seasonally breeding animals often exhibit different social structures during non-breeding and breeding periods that coincide with seasonal environmental variation and resource abundance. However, we know little about the environmental factors associated with when seasonal shifts in social structure occur. This lack of knowledge contrasts with our well-defined knowledge of the environmental cues that trigger a shift to breeding physiology in seasonally breeding species. Here, we identified some of the main environmental factors associated with seasonal shifts in social structure and initiation of breeding in the red-backed fairywren (Malurus melanocephalus), an Australian songbird. Social network analyses revealed that social groups, which are highly territorial during the breeding season, interact in social “communities” on larger home ranges during the non-breeding season. Encounter rates among non-breeding groups were related to photoperiod and rainfall, with shifting photoperiod and increased rainfall associated with a shift toward territorial breeding social structure characterized by reductions in home range size and fewer encounters among non-breeding social groups. Similarly, onset of breeding was highly seasonal and was also associated with non-breeding season rainfall, with greater rainfall leading to earlier breeding. These findings reveal that for some species, the environmental factors associated with the timing of shifts in social structure across seasonal boundaries can be similar to those that determine timing of breeding. This study increases our understanding of the environmental factors associated with seasonal variation in social structure and how the timing of these shifts may respond to changing climates.

Female-limited polymorphisms, where females have multiple forms but males have only one, have been described in a variety of animals, yet are difficult to explain because selection typically is expected to decrease rather than maintain diversity. In the white-necked jacobin ( Florisuga mellivora ), all males and approximately 20% of females express an ornamented plumage type (androchromic), while other females are non-ornamented (heterochromic). Androchrome females benefit from reduced social harassment, but it remains unclear why both morphs persist. Female morphs may represent balanced alternative behavioural strategies, but an alternative hypothesis is that androchrome females are mimicking males. Here, we test a critical prediction of these hypotheses by measuring morphological, physiological and behavioural traits that relate to resource-holding potential (RHP), or competitive ability. In all these traits, we find little difference between female types, but higher RHP in males. These results, together with previous findings in this species, indicate that androchrome females increase access to food resources through mimicry of more aggressive males. Importantly, the mimicry hypothesis provides a clear theoretical pathway for polymorphism maintenance through frequency-dependent selection. Social dominance mimicry, long suspected to operate between species, can therefore also operate within species, leading to polymorphism and perhaps similarities between sexes more generally.

Carotenoid pigments underlie most of the red, orange, and yellow visual signals used in mate choice in vertebrates. However, many of the underlying processes surrounding the production of carotenoid-based traits remain unclear due to the complex nature of carotenoid uptake, metabolism, and deposition across tissues. Here, we leverage the ability to experimentally induce the production of a carotenoid-based red plumage patch in the red-backed fairywren (Malurus melanocephalus), a songbird in which red plumage is an important male sexual signal. We experimentally elevated testosterone in unornamented males lacking red plumage to induce the production of ornamentation and compared gene expression in both the liver and feather follicles between unornamented control males, testosterone-implanted males, and naturally ornamented males. We show that testosterone upregulates the expression of CYP2J19, a gene known to be involved in ketocarotenoid metabolism, and a putative carotenoid processing gene (ELOVL6) in the liver, and also regulates the expression of putative carotenoid transporter genes in red feather follicles on the back, including ABCG1. In black feathers, carotenoid-related genes are downregulated and melanin genes upregulated, but we find that carotenoids are still present in the feathers. This may be due to the activity of the carotenoid-cleaving enzyme BCO2 in black feathers. Our study provides a first working model of a pathway for carotenoid-based trait production in free-living birds, implicates testosterone as a key regulator of carotenoid-associated gene expression, and suggests hormones may coordinate the many processes that underlie the production of these traits across multiple tissues.

Cooperatively breeding species vary widely in degree of social complexity, and disentangling relationships among group members can reveal the costs and benefits of cooperation. Here, we explore the social system of a relatively unstudied cooperatively breeding bird, the Variegated Fairywren (Malurus lamberti), and explore how social complexity and group dynamics may affect cooperation and conflict. We used a combination of field-based population monitoring and detailed social association observations to determine group membership annually across four breeding seasons (2014 – 2017), and used a ddRAD-seq genotyping method to determine genetic relationships within social groups. Social groups ranged in size from 2 – 8 individuals and nearly half of all social groups had multiple adult individuals of both sexes. Approximately two-thirds of those groups exhibited plural breeding, in which multiple females within the same social group nested individually on the same territory. Genetic relationships were diverse across social groups, and many consisted of a combination of relatives and non-relatives of each sex. Notably, although related females often were present within a social group, co-breeding females in the same social group were never closely related to each other. Given extensive variation in relatedness among group members, cooperation in the Variegated Fairywren is likely maintained by a combination of direct and indirect fitness benefits.

During the breeding season, male Red-backed Fairywrens (Malurus melanocephalus) can exhibit ornamented (red-black) or unornamented (brown, resembling females and juveniles) plumage. These distinct plumage types represent alternative reproductive tactics and are associated with behavioural differences during the breeding season. However, we lack an understanding of whether and how these plumage types may be associated with behavioural differences during non-reproductive parts of the year. To fill this knowledge gap, we carried out behavioural observations during the nonbreeding season across three years. We hypothesised that ornamented plumage remains associated with mate attraction behaviours outside of the breeding season. We examined the investment of ornamented, moulting, and unornamented males in social behaviours and found that the three plumage types were largely similar in their behaviour except ornamented males courted and, to a lesser extent, allopreened at higher rates than unornamented males. Since concurrent work in the same study population demonstrates increased extra-pair fitness for males who moult into ornamented plumage early, we speculate that ornamentation and courtship behaviour may serve a mate attraction function outside of the breeding season. We argue that future studies should consider individual-level behavioural monitoring throughout the annual cycle to better quantify the complex selection pressures that lead to the coevolution of plumage moult and alternative reproductive tactics in this system.

Both abiotic environmental conditions and variation in social environment are known to impact the acquisition of sexual signals. However, the influences of abiotic environmental and social factors are rarely compared to each other. Here we test the relative importance of these factors in determining whether and when male red-backed fairy-wrens, Malurus melanocephalus, moult into ornamented breeding plumage, a known sexual signal. One-year-old male red-backed fairy-wrens vary in whether or not they acquire ornamentation, whereas males age 2 years and older vary in their timing of ornament acquisition. It is unclear whether these processes are determined by the same or different factors, and we examined both events using a combination of long-term breeding records and nonbreeding social networks. We found that 1-year-old males that paired prior to the start of the breeding season were more likely to acquire ornamented plumage, but rainfall did not influence whether 1-year-old males acquired ornamented plumage. Thus, for young individuals, social cues appear to play a larger role than abiotic environmental factors in determining ornament acquisition. For older males, timing of ornamented plumage acquisition was constrained by rainfall, with drier nonbreeding seasons leading to poorer physiological condition and later moult dates. Thus, sexual signal variation in older males appears to be a condition-dependent trait, driven by abiotic environmental and physiological factors rather than social cues. These findings reveal that factors influencing sexual signal expression can vary with age when age classes exhibit different forms of signal variation. Our results suggest that social environment may drive sexual signal variation in young individuals, whereas abiotic environmental variation may drive sexual signal variation in older individuals.

Birds are among nature's most social animals. They are renowned for their group migrations, their cooperative foraging, their communal roosting, their synchronous breeding aggregations, their precise parent-offspring interactions, their coordinated group defenses and their intricate territorial and courtship rituals. In these and other contexts, and indeed in most moments of their lives, birds' capacities to navigate complex social demands and relationships can tip the balance between health or sickness, between reproductive success or failure, between life or death. It is thus no surprise that birds have evolved sophisticated systems for mediating social interactions. Foremost among these are communication signals, most evident in birds as songs, calls, color patterns and postural displays. These and other communication signals can transmit information about a signalers' status, motivations and possible future actions, for sensory and cognitive processing by signal receivers. In general, signals provide benefits for both signal senders and receivers. Consider the bright spring plumage of a male wood warbler who has established a breeding territory. The male's plumage pattern benefits him by signaling his 'ownership' of his territory, which can help him attract potential mates and preempt conflict with potential rivals. His plumage pattern also benefits his intended audiences: it informs females about the signaler's potential as a mate, and it informs other males about the degree of threat they might face were they to attempt a territory take-over. Communication signals are, in essence, a glue that maintains the cohesion of bird societies, and a currency that mediates sophisticated relationships within.

During the breeding season, male Red-backed Fairy-wrens Malurus melanocephalus can exhibit a spectrum of plumages from wholly black-and-red to entirely brown. By contrast, females generally appear completely brown, although observations of birds in the hand have revealed that some females also develop some red feathers. Here, we document, for the first time, observations of a female Red-backed Fairy-wren that has developed both black and red feathers under natural conditions. We explore possible explanations for this including whether changes to the individual’s hormonal milieu could be responsible for the unusual plumage that we observed.

Ornamentation, such as the showy plumage of birds, is widespread among female vertebrates, yet the evolutionary pressures shaping female ornamentation remain uncertain. In part this is due to a poor understanding of the mechanistic route to ornamentation in females. To address this issue, we evaluated the evolutionary history of ornament expression in a tropical passerine bird, the White-shouldered Fairywren, whose females, but not males, strongly vary between populations in occurrence of ornamented black-and-white plumage. We first use phylogenomic analysis to demonstrate that female ornamentation is derived and that female ornamentation evolves independently of changes in male plumage. We then use exogenous testosterone in a field experiment to induce partial ornamentation in naturally unornamented females. By sequencing the transcriptome of experimentally induced ornamented and natural feathers, we identify genes expressed during ornament production and evaluate the degree to which female ornamentation in this system is associated with elevated testosterone, as is common in males. We reveal that some ornamentation in females is linked to testosterone and that sexes differ in ornament-linked gene expression. Lastly, using genomic outlier analysis we identify a candidate melanogenesis gene that lies in a region of high genomic divergence among populations that is also differentially expressed in feather follicles of different female plumages. Taken together, these findings are consistent with sex-specific selection favoring the evolution of female ornaments and demonstrate a key role for testosterone in generating population divergence in female ornamentation through gene regulation. More broadly, our work highlights similarities and differences in how ornamentation evolves in the sexes.

The interface between field biology and technology is energizing the collection of vast quantities of environmental data. Passive acoustic monitoring, the use of unattended recording devices to capture environmental sound, is an example where technological advances have facilitated an influx of data that routinely exceeds the capacity for analysis. Computational advances, particularly the integration of machine learning approaches, will support data extraction efforts. However, the analysis and interpretation of these data will require parallel growth in conceptual and technical approaches for data analysis. Here, we use a large hand-annotated dataset to showcase analysis approaches that will become increasingly useful as datasets grow and data extraction can be partially automated.We propose and demonstrate seven technical approaches for analyzing bioacoustic data. These include the following: (1) generating species lists and descriptions of vocal variation, (2) assessing how abiotic factors (e.g., rain and wind) impact vocalization rates, (3) testing for differences in community vocalization activity across sites and habitat types, (4) quantifying the phenology of vocal activity, (5) testing for spatiotemporal correlations in vocalizations within species, (6) among species, and (7) using rarefaction analysis to quantify diversity and optimize bioacoustic sampling.To demonstrate these approaches, we sampled in 2016 and 2018 and used hand annotations of 129,866 bird vocalizations from two forests in New Hampshire, USA, including sites in the Hubbard Brook Experiment Forest where bioacoustic data could be integrated with more than 50 years of observer-based avian studies. Acoustic monitoring revealed differences in community patterns in vocalization activity between forests of different ages, as well as between nearby similar watersheds. Of numerous environmental variables that were evaluated, background noise was most clearly related to vocalization rates. The songbird community included one cluster of species where vocalization rates declined as ambient noise increased and another cluster where vocalization rates declined over the nesting season. In some common species, the number of vocalizations produced per day was correlated at scales of up to 15 km. Rarefaction analyses showed that adding sampling sites increased species detections more than adding sampling days.Although our analyses used hand-annotated data, the methods will extend readily to large-scale automated detection of vocalization events. Such data are likely to become increasingly available as autonomous recording units become more advanced, affordable, and power efficient. Passive acoustic monitoring with human or automated identification at the species level offers growing potential to complement observer-based studies of avian ecology.

Rhodium catalysts for arene alkenylation reported by our group (e.g., Science 2015, 348, 421; J. Am. Chem. Soc. 2017, 139, 5474; J. Am. Chem. Soc. 2018, 140, 17007) have demonstrated selectivity fo...

Directional asymmetry in gonad size is commonly observed in vertebrates and is particularly pronounced in birds, where the left testis is frequently larger than the right. The adaptive significance of directional asymmetry in testis size is poorly understood, and whether it extends beyond the testes (i.e. side-correspondent asymmetry along the reproductive tract) has rarely been considered. Using the Maluridae, a songbird family exhibiting variation in levels of sperm competition and directional testis asymmetry, yet similar in ecology and life history, we investigated the relative roles of side-correspondence and sperm competition on male reproductive tract asymmetry at both inter- and intraspecific levels. We found some evidence for side-correspondent asymmetry. Additionally, sperm competition influenced directional asymmetry at each end of the reproductive tract: species experiencing higher levels of sperm competition had a relatively larger right testis and relatively more sperm in the right seminal glomerus. Within red-backed fairy-wrens (Malurus melanocephalus), auxiliary males had relatively more sperm in the left seminal glomerus, in contrast to a right-bias asymmetry throughout the reproductive tract in breeding males. Given that the number of sperm is important for competitive fertilization success, our results suggest that sperm competition shapes reproductive asymmetries beyond testis size, with likely functional consequences for male reproductive success.

1. Assessing diversity of discretely varying behavior is a classical ethological problem. In particular, the challenge of calculating an individuals' or species' vocal repertoire size is often an important step in ecological and behavioral studies, but a reproducible and broadly applicable method for accomplishing this task is not currently available. 2. We offer a generalizable method to automate the calculation and quantification of acoustic diversity using an unsupervised random forest framework. We tested our method using natural and synthetic datasets of known repertoire sizes that exhibit standardized variation in common acoustic features as well as in recording quality. We tested two approaches to estimate acoustic diversity using the output from unsupervised random forest analyses: (i) cluster analysis to estimate the number of discrete acoustic signals (e.g., repertoire size) and (ii) an estimation of acoustic area in acoustic feature space, as a proxy for repertoire size. 3. We find that our unsupervised analyses classify acoustic structure with high accuracy. Specifically, both approaches accurately estimate element diversity when repertoire size is small to intermediate (5-20 unique elements). However, for larger datasets (20-100 unique elements), we find that calculating the size of the area occupied in acoustic space is a more reliable proxy for estimating repertoire size. 4. We conclude that our implementation of unsupervised random forest analysis offers a generalizable tool that researchers can apply to classify acoustic structure of diverse datasets. Additionally, output from these analyses can be used to compare the distribution and diversity of signals in acoustic space, creating opportunities to quantify and compare the amount of acoustic variation among individuals, populations, or species in a standardized way. We provide R code and examples to aid researchers interested in using these techniques.

Made available in DSpace on 2021-06-25T10:25:43Z (GMT). No. of bitstreams: 0 Previous issue date: 2021-04-01 Animal Behavior Society Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) Temporal coordination of duets consists of nonrandom overlap, alternation or association between rhythms of acoustic elements. Since duet coordination presumably requires high attentiveness between signallers, the coalition quality hypothesis suggests it may indicate the ability or motivation of partners to engage cooperatively in aggressive interactions against intruders. To test this hypothesis, we monitored behavioural as well as vocal responses in a playback experiment conducted with a polyphonal duetting bird, the rufous hornero, Furnarius rufus. We used three categories of treatments in the experiment, which totalled six treatments: duet playbacks that varied in phrase overlap and coordination of temporal rhythms; nonoverlapped (consecutive) solos; and a control heterospecific song. We predicted that birds would respond more strongly to playbacks of coordinated duets than to uncoordinated duets, and also to playbacks of uncoordinated duets than to nonoverlapping male and female solo songs. Coordinated duets (degree of phrase overlap) provoked the weakest response across conspecific stimuli, whereas the responses to uncoordinated duets and nonoverlapped solos were similar. However, partners produced more highly coordinated duets (degree of phrase overlap) in response to coordinated duets (association of phrase rhythms), when compared to all other stimuli. These results suggest that territorial pairs not only perceive but also alter their degree of temporal coordination of duets during territorial interactions. In addition, partners seem to behave cautiously and with more coordinated duets when responding to paired intruders that produce coordinated duets. Our results partially support the coalition quality hypothesis, suggesting that duet coordination may indicate the ability of partners to cooperatively engage in aggressive interactions. Programa de Pós-Graduação em Ecologia Universidade de Brasília Laboratório de Comportamento Animal Departamento de Zoologia Universidade de Brasília Programa de Pós-Graduação em Botânica Universidade de Brasília Laboratório de Fenologia Instituto de Biociências Universidade Estadual Paulista (Unesp) Cornell Lab of Ornithology Department of Neurobiology and Behavior Cornell University Laboratório de Fenologia Instituto de Biociências Universidade Estadual Paulista (Unesp) CNPq: 141178/2015–9 to P.D CNPq: 471945/2013–7 to R.H.M CAPES: 88887.469218/2019–00

Animals produce a wide array of sounds with highly variable acoustic structures. It is possible to understand the causes and consequences of this variation across taxa with phylogenetic comparative analyses. Acoustic and evolutionary analyses are rapidly increasing in sophistication such that choosing appropriate acoustic and evolutionary approaches is increasingly difficult. However, the correct choice of analysis can have profound effects on output and evolutionary inferences. Here, we identify and address some of the challenges for this growing field by providing a roadmap for quantifying and comparing sound in a phylogenetic context for researchers with a broad range of scientific backgrounds. Sound, as a continuous, multidimensional trait can be particularly challenging to measure because it can be hard to identify variables that can be compared across taxa and it is also no small feat to process and analyse the resulting high-dimensional acoustic data using approaches that are appropriate for subsequent evolutionary analysis. Additionally, terminological inconsistencies and the role of learning in the development of acoustic traits need to be considered. Phylogenetic comparative analyses also have their own sets of caveats to consider. We provide a set of recommendations for delimiting acoustic signals into discrete, comparable acoustic units. We also present a three-stage workflow for extracting relevant acoustic data, including options for multivariate analyses and dimensionality reduction that is compatible with phylogenetic comparative analysis. We then summarize available phylogenetic comparative approaches and how they have been used in comparative bioacoustics, and address the limitations of comparative analyses with behavioural data. Lastly, we recommend how to apply these methods to acoustic data across a range of study systems. In this way, we provide an integrated framework to aid in quantitative analysis of cross-taxa variation in animal sounds for comparative phylogenetic analysis. In addition, we advocate the standardization of acoustic terminology across disciplines and taxa, adoption of automated methods for acoustic feature extraction, and establishment of strong data archival practices for acoustic recordings and data analyses. Combining such practices with our proposed workflow will greatly advance the reproducibility, biological interpretation, and longevity of comparative bioacoustic studies.

The existence of distinct traits in island versus mainland populations offers opportunities to gain insights into how eco-evolutionary processes operate under natural conditions. We used two island colonization events in the white-winged fairywren (Malurus leucopterus) to investigate the genomic and demographic origin of melanic plumage. This avian species is distributed across most of Australia, and males of the mainland subspecies (M. l. leuconotus) exhibit a blue nuptial plumage in contrast to males of two island subspecies - M. l. leucopterus on Dirk Hartog Island and M. l. edouardi on Barrow Island - that exhibit a black nuptial plumage. We used reduced-representation sequencing to explore differentiation and demographic history in this species and found clear patterns of divergence between mainland and island populations, with additional substructuring on the mainland. Divergence between the mainland and Dirk Hartog was approximately 10 times more recent than the split between the mainland and Barrow Island, supporting two independent colonizations. In both cases, estimated gene flow between the mainland and the islands was low, contributing to signals of divergence among subspecies. Our results present demographic reconstructions of mainland-island dynamics and associated plumage variation in white-winged fairywrens, with broader implications regarding our understanding of convergent evolution in insular populations.

Specimens have long been viewed as critical to research in the natural sciences because each specimen captures the phenotype (and often the genotype) of a particular individual at a particular point in space and time. In recent years there has been considerable focus on digitizing the many physical specimens currently in the world’s natural history research collections. As a result, a growing number of specimens are each now represented by their own “digital specimen”, that is, a findable, accessible, interoperable and re-usable (FAIR) digital representation of the physical specimen, which contains data about it. At the same time, there has been growing recognition that each digital specimen can be extended, and made more valuable for research, by linking it to data/samples derived from the curated physical specimen itself (e.g., computed tomography (CT) scan imagery, DNA sequences or tissue samples), directly related specimens or data about the organism's life (e.g., specimens of parasites collected from it, photos or recordings of the organism in life, immediate surrounding ecological community), and the wide range of associated specimen-independent data sets and model-based contextualisations (e.g., taxonomic information, conservation status, bioclimatological region, remote sensing images, environmental-climatological data, traditional knowledge, genome annotations). The resulting connected network of extended digital specimens will enable new research on a number of fronts, and indeed this has already begun. The new types of research enabled fall into four distinct but overlapping categories. First, because the digital specimen is a surrogate—acting on the Internet for a physical specimen in a natural science collection—it is amenable to analytical approaches that are simply not possible with physical specimens. For example, digital specimens can serve as training, validation and test sets for predictive process-based or machine learning algorithms, which are opening new doors of discovery and forecasting. Such sophisticated and powerful analytical approaches depend on FAIR, and on extended digital specimen data being as open as possible. These analytical approaches are derived from biodiversity monitoring outputs that are critically needed by the biodiversity community because they are central to conservation efforts at all levels of analysis, from genetics to species to ecosystem diversity. Second, linking specimens to closely associated specimens (potentially across multiple disparate collections) allows for the coordinated co-analysis of those specimens. For example, linking specimens of parasites/pathogens to specimens of the hosts from which they were collected, allows for a powerful new understanding of coevolution, including pathogen range expansion and shifts to new hosts. Similarly, linking specimens of pollinators, their food plants, and their predators can help untangle complex food webs and multi-trophic interactions. Third, linking derived data to their associated voucher specimens increases information richness, density, and robustness, thereby allowing for novel types of analyses, strengthening validation through linked independent data and thus, improving confidence levels and risk assessment. For example, digital representations of specimens, which incorporate e.g., images, CT scans, or vocalizations, may capture important information that otherwise is lost during preservation, such as coloration or behavior. In addition, permanently linking genetic and genomic data to the specimen of the individual from which they were derived—something that is currently done inconsistently—allows for detailed studies of the connections between genotype and phenotype. Furthermore, persistent links to physical specimens, of additional information and associated transactions, are the building blocks of documentation and preservation of chains of custody. The links will also facilitate data cleaning, updating, as well as maintenance of digital specimens and their derived and associated datasets, with ever-expanding research questions and applied uses materializing over time. The resulting high-quality data resources are needed for fact-based decision-making and forecasting based on monitoring, forensics and prediction workflows in conservation, sustainable management and policy-making. Finally, linking specimens to diverse but associated datasets allows for detailed, often transdisciplinary, studies of topics ranging from local adaptation, through the forces driving range expansion and contraction (critically important to our understanding of the consequences of climate change), and social vectors in disease transmission. A network of extended digital specimens will enable new and critically important research and applications in all of these categories, as well as science and uses that we cannot yet envision.

Historically, bird song complexity was thought to evolve primarily through sexual selection on males; yet, in many species, both sexes sing and selection pressure on both sexes may be broader. Previous research suggests competition for mates and resources during short, synchronous breeding seasons leads to more elaborate male songs at high, temperate latitudes. Furthermore, we expect male–female song structure and elaboration to be more similar at lower, tropical latitudes, where longer breeding seasons and year‐round territoriality yield similar social selection pressures in both sexes. However, studies seldom take both types of selective pressures and sexes into account. We examined song in both sexes in 15 populations of nine‐fairy‐wren species (Maluridae), a Southern Hemisphere clade with female song. We compared song elaboration (in both sexes) and sexual song dimorphism to latitude and life‐history variables tied to sexual and social selection pressures and sex roles. Our results suggest that song elaboration evolved in part due to sexual competition in males: male songs were longer than female songs in populations with low male survival and less male provisioning. Also, female songs evolved independently of male songs: female songs were slower paced than male songs, although only in less synchronously breeding populations. We also found male and female songs were more similar when parental care was more equal and when male survival was high, which provides strong evidence that sex role similarity correlates with male–female song similarity. Contrary to Northern Hemisphere latitudinal patterns, male and female songs were more similar at higher, temperate latitudes. These results suggest that selection on song can be sex specific, with male song elaboration favored in contexts with stronger sexual selection. At the same time, selection pressures associated with sex role similarity appear to favor sex role similarity in song structure., Birdsong is often studied as a sexually selected male trait; yet, in many species, both sexes sing. Here, we examined selection pressures responsible for song elaboration and sexual dimorphism in 15 populations of 9 fairy‐wren species (Maluridae), an Australasian lineage with female song. We provide evidence that sex role similarity and sexual selection contribute to male–female song similarity and dimorphism in this group. These results suggest that selection on song can be sex specific, with male song elaboration favored in contexts coincident with sexual selection, while selection pressures associated with sex role similarity also appear to constrain sex‐specific song evolution and song dimorphism.

Natural disturbances like drought and wildfires are expected to increase in prevalence, so understanding how organisms are affected is a key goal for conservationists and biologists alike. While many studies have illustrated long-term effects of perturbations on survival and reproduction, little is known of short-term effects to physiology and sexual signal expression. Ornamental traits have been proposed as reliable indicators of environmental health, yet studies are lacking in the context of natural disturbances. Here we present short-term responses of male Red-backed Fairywrens (Malurus melanocephalus) to wildfire near the onset of the typical breeding season. Males of this species are characterized by plastic expression of sexual plumage phenotypes in their first breeding season. Using two populations with Fairywren captures before and after separate wildfires we illustrate that wildfire suppressed molt into ornamented plumage, including in third year males that typically show little plasticity in ornamentation. Baseline plasma corticosterone was elevated in males sampled after fire, but condition (furcular fat stores) was unaffected. Although testosterone levels did not decrease following fire, we found a positive correlation between testosterone and plumage ornamentation. In addition, males molting in ornamental plumage had higher circulating levels of testosterone than males molting in unornamented plumage following fire. Collectively, these findings suggest that wildfires inhibit or greatly delay acquisition of ornamentation in young males without exerting obvious effects on condition, but rather through subtle effects on testosterone and corticosterone circulation. This natural experiment also reveals that expression of alternative male reproductive phenotypes in this species is sensitive to environmental conditions and more plastic than previously assumed.

Carotenoid pigments produce most red, orange and yellow colours in vertebrates. This coloration can serve as an honest signal of quality that mediates social and mating interactions, but our understanding of the underlying mechanisms that control carotenoid signal production, including how different physiological pathways interact to shape and maintain these signals, remains incomplete. We investigated the role of testosterone in mediating gene expression associated with a red plumage sexual signal in red-backed fairywrens (Malurus melanocephalus). In this species, males within a single population can flexibly produce either red/black nuptial plumage or female-like brown plumage. Combining correlational analyses with a field-based testosterone implant experiment and quantitative polymerase chain reaction, we show that testosterone mediates expression of carotenoid-based plumage in part by regulating expression of CYP2J19, a ketolase gene associated with ketocarotenoid metabolism and pigmentation in birds. This is, to our knowledge, the first time that hormonal regulation of a specific genetic locus has been linked to carotenoid production in a natural context, revealing how endocrine mechanisms produce sexual signals that shape reproductive success.

Sexual selection studies widely estimate several metrics, but values may be inaccurate because standard field methods for studying wild populations produce limited data (e.g., incomplete sampling, inability to observe copulations directly). We compared four selection metrics (Bateman gradient, opportunity for sexual selection, opportunity for selection, and

In socially monogamous species, male reproductive success consists of ‘within-pair’ offspring produced with their socially-paired mate(s), and ‘extra-pair’ offspring produced with additional females throughout the population. Both reproductive pathways offer distinct opportunities for selection in wild populations, as each is composed of separate components of mate attraction, female fecundity, and paternity allocation. Identifying key sources of variance and covariance among these components is a crucial step towards understanding the reproductive strategies that males use to maximize fitness both annually and over their lifetimes. We use 16 years of complete reproductive data from a population of black-throated blue warblers (Setophaga caerulescens) to partition variance in male annual and lifetime reproductive success, and thereby identify if the opportunity for selection varies over the lifetimes of individual males and what reproductive strategies likely favor maximum lifetime fitness. The majority of variance in male reproduction was attributable to within-pair success, but the specific effects of individual components of variance differed between total annual and total lifetime reproductive success. Positive overall lifetime covariance between within-pair and extra-pair components indicates that males able to maximize within-pair success, particularly with double-brooding females, likely achieve higher overall lifetime fitness via both within-pair and extra-pair reproductive pathways.

Polychlorinated biphenyls (PCBs) are worldwide chemical pollutants that have been linked to disrupted reproduction and altered sexual behaviour in many organisms. However, the effect of developmental PCB-exposure on adult passerine reproductive behaviour remains unknown. A commercial PCB mixture (Aroclor 1242) or an estrogenic congener (PCB 52) were administered in sublethal amounts to nestling zebra finches (Taeniopygia guttata) in the laboratory to identify effects of developmental PCB-exposure on adult zebra finch reproductive parameters. Results indicate that although traditional measures of reproductive success are not altered by this PCB dosage, PCBs do alter sexual behaviours such as male song and nesting behaviour. Males treated with PCB 52 in the nest sang significantly fewer syllables than control males, while females treated with Aroclor 1242 in the nest showed the strongest song preferences. PCB treatment also caused an increase in the number of nesting attempts and abandoned nests in the Aroclor 1242 treatment relative to the PCB 52 treatment, and offspring with control fathers fledged significantly earlier than those with fathers treated with Aroclor 1242. Behavioural differences between males seem to best explain these reproductive effects, most notably aggression. These findings suggest that sublethal PCB-exposure during development can significantly alter key reproductive characteristics of adult zebra finches, likely reducing fitness in the wild.

Behavioral changes, such as those involved in mating, foraging, and migration, can generate reproductive barriers between populations. Birds, in particular, are known for their great diversity in these behaviors, and so behavioral isolation is often proposed to be the major driver of speciation. Here, we review empirical evidence to evaluate the importance of behavioral isolation in the early stages of avian speciation. Experimentally measured mating preferences indicate that changes in mating behavior can result in premating barriers, with their strength depending on the extent of divergence in mating signals. Differences in migratory and foraging behavior also can play important roles in generating reproductive barriers in the early stages of speciation. However, because premating behavioral isolation is imperfect, extrinsic postzygotic barriers, in the form of selection against hybrids having intermediate phenotypes, also play an important role in avian diversification, especially in completing the speciation process.

Thermal variation poses a problem for nesting birds and can result in reduced offspring growth rates and survival. To increase the thermal stability of the nest, females can adjust nest characteristics and nest attendance in response to changes in environmental conditions. However, it is unclear how and to what extent females modify parental behaviors during various stages of offspring development. We tested the hypothesis that females adjust nest characteristics and brooding patterns in response to thermal variation during the nest-building and nestling stages, respectively. We examined elevational variation in nest location, nest construction, and brooding patterns in the migratory Black-throated Blue Warbler (Setophaga caerulescens) across a 2°C gradient at the Hubbard Brook Experimental Forest, New Hampshire, USA. Density of woody stems at nest sites and nest wall thickness increased from low to high elevation, corresponding to decreasing temperatures, but we found no relationship between wea...

Information on genetic relationships among individuals is essential to many studies of the behavior and ecology of wild organisms. Parentage and relatedness assays based on large numbers of SNP loci hold substantial advantages over the microsatellite markers traditionally used for these purposes. We present a double-digest restriction site-associated DNA sequencing (ddRAD-seq) analysis pipeline that, as such, simultaneously achieves the SNP discovery and genotyping steps and which is optimized to return a statistically powerful set of SNP markers (typically 150-600 after stringent filtering) from large numbers of individuals (up to 240 per run). We explore the tradeoffs inherent in this approach through a set of experiments in a species with a complex social system, the variegated fairy-wren (Malurus lamberti), and further validate it in a phylogenetically broad set of other bird species. Through direct comparisons with a parallel dataset from a robust panel of highly variable microsatellite markers, we show that this ddRAD-seq approach results in substantially improved power to discriminate among potential relatives and considerably more precise estimates of relatedness coefficients. The pipeline is designed to be universally applicable to all bird species (and with minor modifications to many other taxa), to be cost- and time-efficient, and to be replicable across independent runs such that genotype data from different study periods can be combined and analyzed as field samples are accumulated.

Accurate ageing of nestling birds is important for a variety of ecological, evolutionary and ornithological research. However, this can be difficult when nests are found after the eggs have hatched, especially for less-common species such as brood-parasitic cuckoos. Here we present a photographic guide for ageing nestlings of two species of Australian cuckoo: Horsfield’s Bronze-Cuckoo Chalcites basalis and the Fan-tailed Cuckoo Cacomantis flabelliformis . Evidence of chick rejection by several hosts of bronze-cuckoos Chalcites spp. and counter-adaptations in cuckoo nestlings has prompted increased research attention in this subject. This guide will help facilitate these efforts as well as research into the ecologies of other unstudied cuckoos.

When males compete for mates, they often defend paternity through mate guarding. In addition to physical guarding, in vocal species, especially duetting birds, individuals may duet with their mates in order to guard them. The acoustic mate-guarding hypothesis posits that duetting deters rivals. We experimentally tested the effectiveness of physical and acoustic mate guarding in a duetting songbird, the red-backed fairy-wren, Malurus melanocephalus, using a novel variation of a classic removal experiment. We temporarily removed males, such that females received either no guarding (mate removed), or only acoustic guarding (mate removed, his duet response played from speaker). We found that rival intrusion rates were highest when all guarding was prevented, slightly lower when only acoustic guarding occurred and lowest when pairs were unmanipulated, and both physical and acoustic guarding occurred. This suggests that both guarding techniques deter intruders, but acoustic guarding less so. Intruder display rate during removals was higher than in unmanipulated trials, regardless of acoustic guarding. Results suggest that acoustic guarding may function as a long-range signal that reduces the likelihood of rival intrusion, but physical guarding is necessary to prevent rivals from courting mates. We confirm that physical and acoustic mate guarding serve as important components of intruder deterrence, although they act at different levels. Our study broadens our understanding of multimodal paternity assurance strategies.

Summary Ornamentation is typically observed in sexually mature adults, is often dimorphic in expression, and is most apparent during breeding, supporting a role for sexual selection in its evolution. 1 , 2 , 3 , 4 Yet, increasing evidence suggests that nonsexual social selection may also have a role in the evolution of ornamentation, especially in females. 5 , 6 , 7 , 8 , 9 Distinguishing between these alternatives remains challenging because sexual and nonsexual factors may both play important and overlapping roles in trait evolution. 7 , 10 Here, we show that female ornamentation in a dichromatic hummingbird, the white-necked jacobin (Florisuga mellivora), cannot be explained by sexual selection. Although all males are ornamented, nearly 30% of females have male-like plumage. Remarkably, all juveniles of both sexes express ornamented plumage similar to adult males (androchromatism), but 80% of females acquire non-ornamented plumage (heterochromatism) as they age. This unique ontogeny excludes competition for mates as an explanation for female ornamentation because non-reproductive juveniles are more likely to be ornamented than adults. Instead, avoidance of social harassment appears to underlie this female-limited polymorphism, as heterochrome taxidermy mounts received more aggressive and sexual attention than androchrome mounts from this and other hummingbird species. Monitoring electronically tagged birds at data-logging feeders showed that androchrome females accessed feeders more than heterochrome females, presumably because of reduced harassment. Our findings demonstrate that ornamentation can arise purely through nonsexual social selection, and this hypothesis must be considered in the evolution of not only female-limited polymorphism but also the spectacular ornamentation often assumed to result from sexual selection.

The study of animal acoustic signals is a central tool for many fields in ecology and evolution, but the diversity of analytical methods and sources of animal sound recordings poses important challenges for carrying out robust acoustic analyses. Sound file compression and background noise may both affect acoustic analysis, although little attention has been paid to their respective effects. We evaluated the effect of these factors by assessing the systematic deviation (i.e. bias) and measurement error (i.e. precision) that they generate on spectrographic parameters and two (dis)similarity methods (dynamic time warping on frequency contours and cross-correlation), which represent the most common methods currently used for quantitative characterization of acoustic signals. Measurements were taken across a wide range of signals from a diverse group of bird species, and compared between uncompressed files and decompressed files obtained from mp3-encoded files generated using the two most common mp3 en...

The direct and single-step conversion of benzene, ethylene, and a Cu(II) oxidant to styrene using the Rh(I) catalyst (FlDAB)Rh(TFA)(η2-C2H4) [FlDAB = N,N′-bis(pentafluorophenyl)-2,3-dimethyl-1,4-diaza-1,3-butadiene; TFA = trifluoroacetate] has been reported to give quantitative yields (with Cu(II) as the limiting reagent) and selectivity combined with turnover numbers >800. This report details mechanistic studies of this catalytic process using a combined experimental and computational approach. Examining catalysis with the complex (FlDAB)Rh(OAc)(η2-C2H4) shows that the reaction rate has a dependence on catalyst concentration between first- and half-order that varies with both temperature and ethylene concentration, a first-order dependence on ethylene concentration with saturation at higher concentrations of ethylene, and a zero-order dependence on the concentration of Cu(II) oxidant. The kinetic isotope effect was found to vary linearly with the order in (FlDAB)Rh(OAc)(η2-C2H4), exhibiting no KIE when [...

A series of new rhodium (I) complexes supported by bidentate nitrogen-donor ligands with varying electronic and steric properties were synthesized in situ and evaluated for catalytic arene C−H/D activation. In trifluoroacetic acid (HTFA), these complexes are proposed to mediate H/D exchange of arene C−H/D bonds by an electrophilic aromatic substitution mechanism that involves Rh-mediated activation of HTFA (or DTFA). DFT calculations support the proposed pathway for the H/D exchange reactions.

Duetting has been intensively studied, but we still have little consensus about its fitness consequences. Some studies suggest that duetting functions in acoustic mate guarding to prevent cuckoldry (acoustic paternity guarding hypothesis), whereas other studies argue that duetting is a cooperative behavior to defend common resources (territory defense hypothesis). We tested these 2 hypotheses by investigating the relationships among song traits, extra-pair paternity, territory quality and reproductive success in the Rufous Hornero (Furnarius rufus), a Neotropical, socially monogamous bird. We found a low rate of extra-pair paternity (3.33% of 120 offspring and 6.52% of 46 broods), which suggests that acoustic paternity guarding is probably not a primary function of duetting behavior in this species. Female song output was positively correlated with territory quality, measured as the availability of territory foraging patches. The number of young that survived post-fledging was positively correlated with duet duration, but not with territory features. Our results support the territory defense hypothesis, suggesting that female song in duets might be used in the defense of food resources within territories. Our study provides the first evidence that song traits correlate with both territory features and reproductive success in a duetting species.

In males it is frequently testosterone (T) that activates the expression of sexually selected morphological and behavioral displays, but the role of T in regulating similar traits in females is less clear. Here, we combine correlational data with results from T and gonadotropin-releasing hormone (GnRH) manipulations in both sexes to assess the role of T in mediating sexually dimorphic coloration and morphology in the red-backed fairy-wren (Malurus melanocephalus). We show that: (1) natural variation in female expression of ornamental traits (darkened bills and red back feathers) is positively associated with age and circulating androgen titres, (2) females have the capacity to express most male-typical traits in response to exogenous T, including carotenoid-pigmented body plumage, shorter feathers, darkened bill and enlarged cloacal protuberance, but (3) appear constrained in production of male-typical melanin-pigmented plumage, and (4) low androgen levels during the pre-nuptial molt, probably because of low ovarian capacity for steroid production (or luteinizing hormone sensitivity), prevent females from developing male-like ornamentation. Thus, females appear to retain molecular mechanisms for hormonally regulated male-typical ornamentation, although these are rarely activated because of insufficient production of the hormonal signal.

Single-nucleotide polymorphisms (SNPs) are preferred over microsatellite markers in many evolutionary studies, but have only recently been applied to studies of parentage. Evaluations of SNPs and microsatellites for assigning parentage have mostly focused on special cases that require a relatively large number of heterozygous loci, such as species with low genetic diversity or with complex social structures. We developed 120 SNP markers from a transcriptome assembled using RNA-sequencing of a songbird with the most common avian mating system—social monogamy. We compared the effectiveness of 97 novel SNPs and six previously described microsatellites for assigning paternity in the black-throated blue warbler, Setophaga caerulescens. We show that the full panel of 97 SNPs (mean Ho = 0.19) was as powerful for assigning paternity as the panel of multiallelic microsatellites (mean Ho = 0.86). Paternity assignments using the two marker types were in agreement for 92% of the offspring. Filtering individual samples by a 50% call rate and SNPs by a 75% call rate maximized the number of offspring assigned with 95% confidence using SNPs. We also found that the 40 most heterozygous SNPs (mean Ho = 0.37) had similar power to assign paternity as the full panel of 97 SNPs. These findings demonstrate that a relatively small number of variable SNPs can be effective for parentage analyses in a socially monogamous species. We suggest that the development of SNP markers is advantageous for studies that require high-throughput genotyping or that plan to address a range of ecological and evolutionary questions.