Density dependence is a conceptual cornerstone of avian population biology and, in territorial songbirds, past research has emphasized interactions among food limitation, density, and reproduction. Documenting the importance of density effects is central to understanding how selective forces shape life histories and population dynamics. During the 2008–2011 breeding seasons, we nearly doubled overall conspecific breeding densities on study sites, and manipulated nest box spacing to increase local breeding densities (defined as the number of pairs breeding within 200 m of a pair's nest) of a secondary cavity-nesting songbird, the Prothonotary Warbler (Protonotaria citrea). Our primary objective was to test for effects of food limitation, as mediated by conspecific local densities, on measures of productivity. We monitored breeding pairs and recorded the total number of fledglings produced along with several components of reproductive output (clutch size, hatching success, nestling survival, and probability of attempting a second brood), rates of nestling provisioning, and nestling body condition prior to fledging. We predicted that if the availability of food were affected by local densities, then one or more of these parameters measuring reproduction would be affected negatively. We did not detect an effect of local density on total reproductive output or its components despite our vast range of local densities (1–27 pairs; i.e. 0.16–2.23 pairs ha^–1). Further, we also did not detect differences in nestling provisioning rates and nestling body condition relative to local density. By breeding in a productive ecosystem rich in food resources, these warblers appear to avoid reduced reproductive output when breeding in high densities. Whereas density-dependent food limitation may commonly reduce reproductive output in many species, the ecological circumstances underlying when it does not occur merit further investigation and may provide new insights into what is driving territoriality and what are the primary factors affecting individual fitness.

Tropical mountains feature marked species turnover along elevational gradients and across complex topography, resulting in great concentrations of avian biodiversity. In these landscapes, particularly among morphologically conserved and difficult to observe avian groups, species limits still require clarification. One such lineage is Scytalopus tapaculos, which are among the morphologically most conserved birds. Attention to their distinctive vocal repertoires and phylogenetic relationships has resulted in a proliferation of newly identified species, many of which are restricted range endemics. Here, we present a revised taxonomy and identify species limits among high-elevation populations of Scytalopus tapaculos inhabiting the Peruvian Andes. We employ an integrated framework using a combination of vocal information, mitochondrial DNA sequences, and appearance, gathered from our own fieldwork over the past 40 yr and supplemented with community-shared birdsong archives and museum specimens. We describe 3 new species endemic to Peru. Within all 3 of these species there is genetic differentiation, which in 2 species is mirrored by subtle geographic plumage and vocal variation. In a fourth species, Scytalopus schulenbergi, we document deep genetic divergence and plumage differences despite overall vocal similarity. We further propose that an extralimital taxon, Scytalopus opacus androstictus, be elevated to species rank, based on a diagnostic vocal character. Our results demonstrate that basic exploration and descriptive work using diverse data sources continues to identify new species of birds, particularly in tropical environs.

Optimizing breeding phenology, an important aspect of fitness, is complex for migratory species as they must make key timing decisions early, and remotely, from breeding sites. We examined the role of weather (locally and cross-seasonally), cavity availability, and competitive exclusion in determining among-year variation in breeding phenology over 17 yr for 2 migratory, cavity-nesting birds: Mountain Bluebirds (Sialia currucoides; n = 462 nests) and Tree Swallows (Tachycineta bicolor; n = 572) using natural tree cavities in British Columbia, Canada. We assessed weather effects within the winter and migratory range and at our study sites. We quantified competition as the proportion of cavities occupied by European Starlings (Sturnus vulgaris) (for both species) and Mountain Bluebirds (for Tree Swallow only) in each year. For 229 bluebird and 177 swallow nests with known fates, we tested whether late years resulted in reduced productivity. Although the effects were small, heavy rainfall and strong diurnal westerly winds during migration were associated with breeding delays for Mountain Bluebirds. However, cavity availability (earlier breeding with increases) had a 5–8 × greater effect on timing than migratory conditions. There was no evidence that starling competition delayed bluebirds. In Tree Swallows, greater local daily rainfall was associated with delayed breeding, as was starling abundance (the effect of starlings was 1.4 × smaller than that of rainfall). Neither bluebird abundance nor cavity availability changed swallow phenology. Neither species showed reduced productivity in late breeding years. In both species, individuals that bred late relative to conspecifics within-year had smaller clutches and greater probability of nest failure. We conclude that breeding ground conditions, particularly cavity limitation and local rainfall (for swallows), are important drivers of breeding phenology for our focal species, but that the productivity cost of late years, at least for Tree Swallows, is minimal.

Natal dispersal and local recruitment are affected by factors both intrinsic and extrinsic to juveniles and may affect fitness. Understanding the relationship between dispersal and population density in birds has been hindered by a lack of long-term studies and a focus on resident species has neglected the role of weather operating at large spatial scales. I studied local recruitment and the reproductive consequences of natal dispersal distance within a population of Northern Flickers (Colaptes auratus), a migratory woodpecker. During a field study spanning 16 yr in British Columbia, 8,272 fledglings were banded and 138 males and 105 females recruited locally. The average annual local recruitment rate for males (3.36%) was greater than that for females (2.55%) and the propensity to recruit locally was positively correlated with an early hatch date and high body condition. Annual local recruitment was not associated with population density in the year of hatch but was positively correlated with population density in the year of settlement. Local recruitment was also positively correlated with warmer springs during migration, consistent with the phenology hypothesis that the location of settlement is affected by weather along the route. Among local recruits, natal dispersal distance was independent of the presence of parents and so dispersal was not a behavior to prevent inbreeding. However, settling closer to the natal site led to reproductive benefits in terms of earlier laying dates and better nest success. Therefore, juveniles may gain useful information about the location of nesting substrates, predation risk, and patchy food resources by exploring the landscape around their natal site during the post-fledging period and then settling in the familiar area after returning from migration.

Migratory birds are demonstrating changes in phenology linked to climate change. Understanding these changes requires connecting events that occur over the multiple regions occupied during their annual cycle. The Burrowing Owl (Athene cunicularia) is a species of concern in North America, with pronounced declines in regions of the Great Plains. Using a dataset that spanned 10 breeding sites from South Dakota to northern Mexico in various years during 1989–2017, we observed both advances and delays in nesting along with increasing variation in nest initiation dates. We examined the effects of a large-scale climate system (El Niño Southern Oscillation), drought, and local weather patterns throughout the annual cycle as potential predictors of early and late nesting. Moisture conditions during the winter and spring migratory period had the greatest influence on nest phenology. Years with more intense drought on winter and migratory grounds increased the probability of nests initiating late relative to early. Correspondingly, wet conditions were associated with an increased probability of early nest initiation. Drought likely has cascading ecological effects that negatively influence food abundance for Burrowing Owls, resulting in delays in the ability of individuals to meet energetic demands required for migration. How climate change will impact Burrowing Owl phenology is important considering a projected increase in the magnitude and frequency of drought and declining owl population trends.

Grooming by birds is thought to serve essential anti-parasite functions. While preening has been well studied, little is known about the function of scratching in birds. We conducted a series of experiments to determine the effectiveness of scratching for controlling feather lice (Columbicola columbae) on Rock Pigeons (Columba livia). First, we used a hobbling technique to impair scratching. After 6 mo, hobbled birds had significantly more lice than controls that could scratch. In addition, lice on hobbled birds were concentrated on the birds' heads and necks (i.e. the regions that birds scratch). Secondly, we tested the role the claw plays in scratching by declawing nestlings. Once mature, declawed pigeons had significantly more lice than control birds with claws. Moreover, lice on declawed birds were concentrated on the head and neck. Next, we tested whether the flange found on the middle claw of many bird species enhances scratching. We experimentally manipulated the flange; however, the number and location of lice on birds without flanges was not significantly different than that on control birds with intact flanges. Finally, we tested whether scratching removes parasites directly or indirectly by “flushing” them onto body regions where they can be preened. When we impaired scratching (with hobbles) and preening (with “bits”) we found that scratching no longer reduced the number of lice on birds. Our results indicated that scratching and preening work synergistically; scratching reduces parasite load by flushing lice onto regions of the body where they can be eliminated by preening.

We studied the phylogeny, biogeography, and diversification of suboscine passerines in the genus Scytalopus (Rhinocryptidae), a widespread, species-rich, and taxonomically challenging group of Neotropical birds. We analyzed nuclear (exons, regions flanking ultraconserved elements) and mitochondrial (ND2) DNA sequence data for a taxonomically and geographically comprehensive sample of specimens collected from Costa Rica to Patagonia and Brazil. We found that Scytalopus is a monophyletic group sister to Eugralla and consists of 3 main clades roughly distributed in (1) the Southern Andes, (2) eastern Brazil, and (3) the Tropical Andes and Central America. The clades from the Southern Andes and eastern Brazil are sister to each other. Despite their confusing uniformity in plumage coloration, body shape, and overall appearance, rates of species accumulation through time in Scytalopus since the origin of the clade in the Late Miocene are unusually high compared with those of other birds, suggesting rapid non-adaptive diversification in the group. We attribute this to their limited dispersal abilities making them speciation-prone and their occurrence in a complex landscape with numerous barriers promoting allopatric differentiation. Divergence times among species and downturns in species accumulation rates in recent times suggest that most speciation events in Scytalopus predate climatic oscillations of the Pleistocene. Our analyses identified various cases of strong genetic structure within species and lack of monophyly of taxa, flagging populations which likely merit additional study to clarify their taxonomic status. In particular, detailed analyses of species limits are due in S. parvirostris, S. latrans, S. speluncae, the S. atratus complex, and the Southern Andes clade.

In all animals, susceptibility to parasites can differ among individuals. Young, nest-bound birds are exposed to a diversity of nest-dwelling ectoparasites that typically feed on their blood. Within broods, hatching asynchrony creates size hierarchies that result in morphological and physiological variation among nest mates, and susceptibility to parasites also may vary predictably with this size hierarchy. Our objective was to use a broad-spectrum, anti-parasite drug, ivermectin (IVM), to treat individual nestling Tree Swallows (Tachycineta bicolor) and assess how nestling susceptibility to parasites varied both within and among broods. Broods were either assigned to an IVM group, where half of the nestlings in a brood received IVM injections and half received control injections of pure sesame oil, or to a control group, where all nestlings received oil injections. We found that the IVM treatment reduced parasite loads for broods as a whole, thereby benefiting all nestlings in IVM broods and suggesting our treatment resulted in herd immunity. Specifically, nestlings from IVM broods had higher hemoglobin concentrations, regardless of whether they received injections with IVM or oil, and greater fledging success, than nestlings from control broods. On the contrary, IVM treatment did not strongly affect nestling morphology, with only marginal effects on the growth rate of ninth primary feathers, and the effects of the treatment on 2 other morphological traits depending on temporal factors. Variation in size within broods, however, influenced the chance of an individual fledging, which increased with relative size within a brood, but only under lower parasite loads (i.e. IVM broods). By experimentally manipulating nestling susceptibility to parasites, we have demonstrated variation in nestling response to an anti-parasite treatment both within and among broods, and future studies should investigate the underlying mechanism for why certain nestlings along the brood size hierarchy are more susceptible to parasites.

Mixed-species flocks are ubiquitous in forest bird communities, yet the extent to which positive (facilitative) or negative (competitive) interactions structure these assemblages has been a subject of debate. Here, we describe the fine-scale foraging ecology and use network analysis to quantify mixed-species flocking interactions of an insectivorous bird community in hardwood forests of north-central Florida. Our goal was to determine if similarly foraging species are more (facilitation hypothesis) or less (competition hypothesis) likely to associate in flocks, and if foraging ecology can explain intraspecific abundance patterns within flocks. We quantified attack maneuvers, foraging substrate, and foraging microhabitat of all 17 common insectivorous species in these forests and characterized the composition of 92 flocks encountered. Flocking was important in our community; 14 of 17 species joined more than 5% of flocks, and 10 species had flocking propensities of over 0.80. Our results supported both hypothesized mechanisms structuring flock composition. Species had distinct, well-defined foraging niches during the nonbreeding season, but foraging niche overlap among flocking species was greater than expected by chance. Consistent with the facilitation hypothesis, we found that similarly foraging species were significantly more likely to associate in flocks, a result driven by lower association strengths in large-bodied woodpeckers. We found no evidence of assortment by foraging behavior, however, likely because foraging behavior and substrate use showed strong niche partitioning at the fine scale within our community. Intraspecific abundance patterns were significantly linked to foraging substrate use, with live leaf use correlated with high within-flock abundance and relative abundance at study sites. Species that specialized on comparatively less abundant substrates (tree trunks, epiphytes, dead leaves) joined flocks as singletons, showed lower relative abundance, and may exhibit nonbreeding territoriality. Our results highlight the importance of foraging substrate use and mixed-species flocks in structuring the nonbreeding ecology of migratory birds.

The availability of detailed information that encompasses the geographic range of a species, spans a long-term temporal range, and yields individual information (e.g., age and sex), is a principle challenge in ecology. To this end, the North American Bird Banding Laboratory maintains a unique and underutilized dataset that can be used to address core questions of phenological change in migratory birds. We used records from 1966 to 2015 to quantify how the timing of migration has shifted in a long-distance migrant, the Black-throated Blue Warbler (Setophaga caerulescens). Additionally, we examined age and sex differences in the timing of migratory movements. We observed that early spring migrants passed through sites ∼1.1 days earlier per decade and the peak of spring migration also occurred earlier over the 50 yr of this study. Additionally, phenological change was more rapid with increasing latitude during peak spring migratory periods. During fall, the peak of migration stayed consistent across the 50 yr studied, but the migratory season showed protraction overall. During spring, males consistently migrated earlier than females and adults migrated earlier than young individuals. During fall, there was no difference in timing between males and females, but young birds migrated earlier than adults. Additionally, migration proceeded faster in spring compared with the fall. This study reveals differential strategies in migrant timing, across seasons, age groups, and by sex, and shows that en route adjustments across latitude may account for changes in migrant timing. This basic information about such a fundamental ecological process is crucial to our understanding of migration and we must utilize these unique data to appreciate critical shifts at relevant scales of migration.