A late burst of colour evolution in a radiation of songbirds (Passeriformes: Parulidae) suggests secondary contact drives signal divergence.

Evolutionary radiations provide important insights into species diversification, which is especially true of adaptive radiations. New World wood warblers (Parulidae) are a family of small, insectivorous, forest-dwelling passerine birds, often considered an exemplar of adaptive radiation due to their rapid diversification followed by a slowdown. However, they deviate from the expectations of an adaptive radiation scenario due to the lack of conspicuous morphological and ecological differentiation. We fitted several macroevolutionary models to trait data in 105 species of wood warblers. We tested whether morphological traits underwent an early burst of evolution (suggesting adaptation to new ecological niches in adaptive radiations) and whether song and colour underwent a diversity-dependent acceleration of trait evolutionary rate (consistent with reproductive interference driving signal evolution). Morphology and song evolved gradually under stabilizing selection, suggesting niche conservatism, with morphology possibly acting as a constraint on song evolution. In contrast, many feather colour traits underwent a diversity-dependent burst of evolution occurring late in the clade's history. We suggest that a two-step process has led to the remarkable diversification of wood warblers. First, their early diversification probably proceeded by allopatric speciation. Second, feather colour divergence likely occurred during secondary contact after range expansion. This diversification of signalling traits might have facilitated species coexistence, in combination with behavioural niche partitioning. Wood warblers seem to present characteristics of both adaptive and non-adaptive radiations.

AVONET: morphological, ecological and geographical data for all birds.

Functional traits offer a rich quantitative framework for developing and testing theories in evolutionary biology, ecology and ecosystem science. However, the potential of functional traits to drive theoretical advances and refine models of global change can only be fully realised when species-level information is complete. Here we present the AVONET dataset containing comprehensive functional trait data for all birds, including six ecological variables, 11 continuous morphological traits, and information on range size and location. Raw morphological measurements are presented from 90,020 individuals of 11,009 extant bird species sampled from 181 countries. These data are also summarised as species averages in three taxonomic formats, allowing integration with a global phylogeny, geographical range maps, IUCN Red List data and the eBird citizen science database. The AVONET dataset provides the most detailed picture of continuous trait variation for any major radiation of organisms, offering a global template for testing hypotheses and exploring the evolutionary origins, structure and functioning of biodiversity.

Functional diversity of avian communities increases with canopy height: From individual behavior to continental-scale patterns.

Vegetation complexity is an important predictor of animal species diversity. Specifically, taller vegetation should provide more potential ecological niches and thus harbor communities with higher species richness and functional diversity (FD). Resource use behavior is an especially important functional trait because it links species to their resource base with direct relevance to niche partitioning. However, it is unclear how exactly the diversity of resource use behavior changes with vegetation complexity. To address this question, we studied avian FD in relation to vegetation complexity along a continental-scale vegetation gradient. We quantified foraging behavior of passerine birds in terms of foraging method and substrate use at 21 sites (63 transects) spanning 3,000 km of woodlands and forests in Australia. We also quantified vegetation structure on 630 sampling points at the same sites. Additionally, we measured morphological traits for all 111 observed species in museum collections. We calculated individual-based, abundance-weighted FD in morphology and foraging behavior and related it to species richness and vegetation complexity (indexed by canopy height) using structural equation modeling, rarefaction analyses, and distance-based metrics. FD of morphology and foraging methods was best predicted by species richness. However, FD of substrate use was best predicted by canopy height (ranging 10-30 m), but only when substrates were categorized with fine resolution (17 categories), not when categorized coarsely (8 categories). These results suggest that, first, FD might increase with vegetation complexity independently of species richness, but whether it does so depends on the studied functional trait. Second, patterns found might be shaped by how finely we categorize functional traits. More complex vegetation provided larger "ecological space" with more resources, allowing the coexistence of more species with disproportionately more diverse foraging substrate use. We suggest that the latter pattern was driven by nonrandom accumulation of functionally distinct species with increasing canopy height.

Adaptation and constraint shape the evolution of growth patterns in passerine birds across the globe.

BACKGROUND: Growth trajectories should be adapted to selective factors of each species' environment. However, major shaping forces of growth and development are unclear, especially when studying several traits at once. Birds provide an ideal opportunity to analyze growth patterns across species due to there being enough available data. We tested the relative importance of nest predation risk, the number of care-givers, nest height, foraging substrate, clutch size, and latitude on growth patterns of passerine birds (Passeriformes) using phylogenetic comparative methods. Specifically, we studied the evolution of fledging time, average and peak growth rates, and relative development at fledging of body mass and tarsus, wing, and tail length. RESULTS: Using a comprehensive literature search and data quality control, we obtained data on growth in 231 species based on 295 populations. Species with long development in the nest grew slowly and had well-developed traits at fledging. Species breeding under high nest predation risk, building their nests close to the ground, and those living in northern temperate regions fledged early and grew fast, sometimes fledging with less developed body mass and traits critical for locomotion (tarsus, wing, and tail). On the other hand, the number of caring adults, clutch size, and species' foraging substrate had very limited predictive value for growth patterns across passerine species. CONCLUSIONS: Shortening of the nestling period was a primary means of accelerating development (in relation to nest predation, nest height, and latitude), sometimes supplemented by higher peak growth rates of body mass, tarsus, and wing (especially in relation to latitude). Overall growth patterns of passerines were adaptively tuned to nest predation risk and nest height, with northern temperate species having especially short nestling periods and fast growth rates of body mass, tarsus, and wing.

Evolution of parental activity at the nest is shaped by the risk of nest predation and ambient temperature across bird species.

Incubation is an important component of parental care in birds, and species differ widely in their incubation rhythm. In this comparative study, we focused on factors responsible for those differences. As hypothesized by A. Skutch, increased parental activity at the nest increases the probability of nest depredation. High risk of nest predation should therefore lead to the evolution of lower frequency of parental activity at the nest. We thus expected to find a negative relationship between frequency of nest visits and the risk of nest depredation. Using a large dataset of 256 species of passerines breeding worldwide, we found that the frequency of nest visits decreased as the risk of nest depredation increased and that this effect was strongest in tropical species. Further, foraging bouts were longer in species experiencing warmer ambient temperatures during incubation and those with domed nests. Incubation bouts were longer and frequency of nest visits was lower in species with higher body mass. Our results support the view that natural selection favors lower frequency of nests visits in species under higher risk of nest predation and demonstrate the importance of other factors (temperature, geographic space, nest type, and body mass) in shaping the evolution of incubation rhythm.

The importance of having a partner: male help releases females from time limitation during incubation in birds.

INTRODUCTION: Male contribution to parental care varies widely among avian species. Yet the reasons for this variation, as well as its consequences, are still unclear. Because the amount of care provided by one sex is ultimately constrained by the time available for energy acquisition, contribution by the other sex should increase when overall parental workload is high. We tested this prediction by analyzing male contribution to incubation in 528 populations of 320 species of passerines, where females usually devote more time to incubation than males. Our worldwide sample included species with female-only parental care (the male is not present), incubation feeding (the male feeds the incubating female), and shared incubation (both sexes incubate the eggs). RESULTS: Overall nest attentiveness was greatest in species with shared incubation followed by species with incubation feeding and lowest in species with female-only care. Nest attentiveness and the degree of male contribution to incubation in species with shared incubation were very strongly correlated, whereas this correlation was absent in females. Interestingly, female contribution decreased towards the equator while male contribution did not change significantly with latitude. Hence, relative male incubation effort increased towards the equator, whereas that of female decreased. In species with incubation feeding, female nest attentiveness increased with the frequency of male feeding. CONCLUSIONS: These findings support the hypothesis that male help is indispensable for increasing nest attentiveness in birds, either in the form of incubation feeding (supply of energy) or direct incubation of eggs. We conclude that energy acquisition constraints might be a potent force driving sex-specific contribution to parental care.

More ornamented females produce higher-quality offspring in a socially monogamous bird: an experimental study in the great tit (Parus major).

INTRODUCTION: Animals are often conspicuously colored and explanations range from aposematism and mimicry to sexual selection. Although sexual selection explains vivid coloration in males, functional significance of vivid coloration in females of socially monogamous species remains unclear. The hypothesis of mutual mate choice predicts that more ornamented females produce offspring of higher quality. We tested this prediction in the great tit (Parus major), a small, insectivorous, socially monogamous passerine. RESULTS: In both females and males we quantified three ornaments that have been hypothesized to have signaling role in this species (size of black breast stripe, carotenoid chroma of yellow breast feathers, immaculateness of the white cheek). We swapped broods between nests soon after hatching, thus separating genetic plus pre-hatching vs. post-hatching effects on offspring performance. Body mass of offspring at 14 days of age was positively related to the area of black breast stripe of genetic mothers. Immune response to a novel antigen (phytohaemagglutinin) at 14 days of age was positively related to the immaculateness of the white cheek patch of both genetic and foster mothers. CONCLUSIONS: We showed that females with more elaborate ornaments produced higher-quality offspring and we discuss potential proximate mechanisms of these relationships. We conclude that as more elaborate ornaments were reliable signals of offspring quality, direct selection by male mate choice might have been responsible for the evolution and/or maintenance of these signaling traits in females.