Complex patterns of morphological diversity across multiple populations of an urban bird species.

Urbanization presents a natural evolutionary experiment because selection pressures in cities can be strongly mismatched with those found in species' historic habitats. However, some species have managed to adapt and even thrive in these novel conditions. When a species persists across multiple cities, a fundamental question arises: do we see similar traits evolve under similar novel environments? By testing if and how similar phenotypes emerge across multiple urban populations, we can begin to assess the predictability of population response to anthropogenic change. Here, we examine variation within and across multiple populations of a songbird, the dark-eyed junco (Junco hyemalis). We measured morphological variations in juncos across urban and non-urban populations in Southern California. We investigated whether the variations we observed are due to differences in environmental conditions across cities. Bill shape differed across urban populations; Los Angeles and Santa Barbara juncos had shorter, deeper bills than non-urban juncos, but San Diego juncos did not. On the other hand, wing length decreased with the built environment, regardless of the population. Southern Californian urban juncos exhibit both similarities and differences in morphological traits. Studying multiple urban populations can help us determine the predictability of phenotypic evolutionary response to novel environments.

Urban birds become less fearful following COVID-19 reopenings.

Following the COVID-19 pandemic, many people around the world stayed home, drastically altering human activity in cities. This exceptional moment provided researchers the opportunity to test how urban animals respond to human disturbance, in some cases testing fundamental questions on the mechanistic impact of urban behaviours on animal behaviour. However, at the end of this 'anthropause', human activity returned to cities. How might each of these strong shifts affect wildlife in the short and long term? We focused on fear response, a trait essential to tolerating urban life. We measured flight initiation distance-at both individual and population levels-for an urban bird before, during and after the anthropause to examine if birds experienced longer-term changes after a year and a half of lowered human presence. Dark-eyed juncos did not change fear levels during the anthropause, but they became drastically less fearful afterwards. These surprising and counterintuitive findings, made possible by following the behaviour of individuals over time, has led to a novel understanding that fear response can be driven by plasticity, yet not habituation-like processes. The pandemic-caused changes in human activity have shown that there is great complexity in how humans modify a behavioural trait fundamental to urban tolerance in animals.

Meta-analysis of three-stressor combinations on population-level fitness reveal substantial higher-order interactions.

Although natural populations are typically subjected to multiple stressors, most past research has focused on single-stressor and two-stressor interactions, with little attention paid to higher-order interactions among three or more stressors. However, higher-order interactions increasingly appear to be widespread. Consequently, we used a recently introduced and improved framework to re-analyze higher-order ecological interactions. We conducted a literature review of the last 100 years (1920-2020) and reanalyzed 142 ecological three-stressor interactions on species' populations from 38 published papers; the vast majority of these studies were from the past 10 years. We found that 95.8 % (n = 136) of the three-stressor combinations had either not been categorized before or resulted in different interactions than previously reported. We also found substantial levels of emergent properties-interactions that are not due to strong pairwise interactions within the combination but rather uniquely due to all three stressors being combined. Calculating net interactions-the overall accounting for all possible interactions within a combination including the emergent and all pairwise interactions-we found that the most prevalent interaction type is antagonism, corresponding to a smaller than expected effect based on single stressor effects. In contrast, for emergent interactions, the most prevalent interaction type is synergistic, resulting in a larger than expected effect based on single stressor effects. Additionally, we found that hidden suppressive interactions-where a pairwise interaction is suppressed by a third stressor-are found in the majority of combinations (74 %). Collectively, understanding multiple stressor interactions through applying an appropriate framework is crucial for answering fundamental questions in ecology and has implications for conservation biology and population management. Crucially, identifying emergent properties can reveal hidden suppressive interactions that could be particularly important for the ecological management of at-risk populations.

No evidence of repeated song divergence across multiple urban and non-urban populations of dark-eyed juncos (Junco hyemalis) in Southern California.

Urbanization can affect species communication by introducing new selection pressures, such as noise pollution and different environmental transmission properties. These selection pressures can trigger divergence between urban and non-urban populations. Songbirds rely on vocalizations to defend territories and attract mates. Urban songbirds have been shown in some species and some populations to increase the frequencies, reduce the length and change other temporal features of their songs. This study compares songs from four urban and three non-urban populations of dark-eyed juncos (Junco hyemalis) throughout Southern California. We examined song length, trill rate, minimum frequency, maximum frequency, peak frequency and frequency bandwidth. We also compared songs recorded from one urban junco population in San Diego nearly two decades ago with recently collected data in 2018-2020. Over all comparisons, we only found significant differences between UCLA and the 2006/2007 UCSD field seasons in minimum and maximum frequencies. These findings partially support and are partially in contrast to previous urban song studies. As urban areas expand, more opportunities will arise to understand urban song divergence in greater detail.

Male-like female morphs in hummingbirds: the evolution of a widespread sex-limited plumage polymorphism.

Differences in the way males and females look or behave are common in animals. However, discrete variation within sexes (sex-limited polymorphism) also occurs in several vertebrate and invertebrate lineages. In birds, female-limited polymorphism (FLP) in which some females resemble males in coloration is most prominent in hummingbirds, a group known for its morphological and behavioural sexual dimorphism. Yet, it remains unclear whether this intrasexual colour variation in hummingbirds arises through direct selection on females, or indirectly as a non-adaptive byproduct resulting from selection on males. Here, we analysed specimens from more than 300 hummingbird species to determine the extent, evolutionary history and function of FLP. We found that FLP evolved independently in every major clade and occurs in nearly 25% of hummingbird species. Using phylogenetically informed analyses, we rejected non-adaptive hypotheses that FLP is the result of indirect selection or pleiotropy across species. Instead, FLP is associated with ecology, migratory status, and marginally with social dominance, suggesting a socioecological benefit to females. Ultimately, we show that FLP is not only widespread in hummingbirds and likely adaptive, but may also be useful for understanding the evolution of female ornamentation in systems under strong sexual selection.

Nests in the cities: adaptive and non-adaptive phenotypic plasticity and convergence in an urban bird.

Phenotypic plasticity plays a critical role in adaptation to novel environments. Behavioural plasticity enables more rapid responses to unfamiliar conditions than evolution by natural selection. Urban ecosystems are one such novel environment in which behavioural plasticity has been documented. However, whether such plasticity is adaptive, and if plasticity is convergent among urban populations, is poorly understood. We studied the nesting biology of an 'urban-adapter' species, the dark-eyed junco (Junco hyemalis), to understand the role of plasticity in adapting to city life. We examined (i) whether novel nesting behaviours are adaptive, (ii) whether pairs modify nest characteristics in response to prior outcomes, and (iii) whether two urban populations exhibit similar nesting behaviour. We monitored 170 junco nests in urban Los Angeles and compared our results with prior research on 579 nests from urban San Diego. We found that nests placed in ecologically novel locations (off-ground and on artificial surfaces) increased fitness, and that pairs practiced informed re-nesting in site selection. The Los Angeles population more frequently nested off-ground than the San Diego population and exhibited a higher success rate. Our findings suggest that plasticity facilitates adaptation to urban environments, and that the drivers behind novel nesting behaviours are complex and multifaceted.

Using a newly introduced framework to measure ecological stressor interactions.

Understanding how stressors combine to affect population abundances and trajectories is a fundamental ecological problem with increasingly important implications worldwide. Generalisations about interactions among stressors are challenging due to different categorisation methods and how stressors vary across species and systems. Here, we propose using a newly introduced framework to analyse data from the last 25 years on ecological stressor interactions, for example combined effects of temperature, salinity and nutrients on population survival and growth. We contrast our results with the most commonly used existing method - analysis of variance (ANOVA) - and show that ANOVA assumptions are often violated and have inherent limitations for detecting interactions. Moreover, we argue that rescaling - examining relative rather than absolute responses - is critical for ensuring that any interaction measure is independent of the strength of single-stressor effects. In contrast, non-rescaled measures - like ANOVA - find fewer interactions when single-stressor effects are weak. After re-examining 840 two-stressor combinations, we conclude that antagonism and additivity are the most frequent interaction types, in strong contrast to previous reports that synergy dominates yet supportive of more recent studies that find more antagonism. Consequently, measuring and re-assessing the frequency of stressor interaction types is imperative for a better understanding of how stressors affect populations.