Host space, not energy or symbiont size, constrains feather mite abundance across passerine bird species.

Comprehending symbiont abundance among host species is a major ecological endeavour, and the metabolic theory of ecology has been proposed to understand what constrains symbiont populations. We parameterized metabolic theory equations to investigate how bird species' body size and the body size of their feather mites relate to mite abundance according to four potential energy (uropygial gland size) and space constraints (wing area, total length of barbs and number of feather barbs). Predictions were compared with the empirical scaling of feather mite abundance across 106 passerine bird species (26,604 individual birds sampled), using phylogenetic modelling and quantile regression. Feather mite abundance was strongly constrained by host space (number of feather barbs) but not by energy. Moreover, feather mite species' body size was unrelated to the body size of their host species. We discuss the implications of our results for our understanding of the bird-feather mite system and for symbiont abundance in general.

Rethinking megafauna.

Concern for megafauna is increasing among scientists and non-scientists. Many studies have emphasized that megafauna play prominent ecological roles and provide important ecosystem services to humanity. But, what precisely are 'megafauna'? Here, we critically assess the concept of megafauna and propose a goal-oriented framework for megafaunal research. First, we review definitions of megafauna and analyse associated terminology in the scientific literature. Second, we conduct a survey among ecologists and palaeontologists to assess the species traits used to identify and define megafauna. Our review indicates that definitions are highly dependent on the study ecosystem and research question, and primarily rely on ad hoc size-related criteria. Our survey suggests that body size is crucial, but not necessarily sufficient, for addressing the different applications of the term megafauna. Thus, after discussing the pros and cons of existing definitions, we propose an additional approach by defining two function-oriented megafaunal concepts: 'keystone megafauna' and 'functional megafauna', with its variant 'apex megafauna'. Assessing megafauna from a functional perspective could challenge the perception that there may not be a unifying definition of megafauna that can be applied to all eco-evolutionary narratives. In addition, using functional definitions of megafauna could be especially conducive to cross-disciplinary understanding and cooperation, improvement of conservation policy and practice, and strengthening of public perception. As megafaunal research advances, we encourage scientists to unambiguously define how they use the term 'megafauna' and to present the logic underpinning their definition.

Genes acting in synapses and neuron projections are early targets of selection during urban colonization.

When a species colonizes an urban habitat, differences in the environment can create novel selection pressures. Successful colonization will further lead to demographic perturbations and genetic drift, which can interfere with selection. Here, we test for consistent urban selection signals in multiple populations of the burrowing owl (Athene cunicularia), a species that colonized South American cities just a few decades ago. We sequenced 213 owls from three urban-rural population pairs and performed a genome-wide comparison of urban against rural birds. We further studied genome-wide associations with flight initiation distance, a measure of harm avoidance in which urban and rural birds are known to differ. Based on four samples taken over nine years from one of the urban populations, we investigated temporal allele frequency changes. The genomic data were also used to identify urban-specific signatures of selective sweeps. Single genomic sites did not reach genome-wide significance for any association. However, a gene-set analysis on the strongest signals from these four selection scans suggests a significant enrichment of genes with known functions related to synapses and neuron projections. We identified 98 genes predominantly expressed in the brain, of which many may play a role in the modulation of brain connectivity and consequently in cognitive function and motivational behaviour during urbanization. Furthermore, polymorphisms in the promoter region of the synaptic SERT gene - one of the two candidates known to correlate with urban colonization in birds - associated with the habitat in which individuals lived (urban vs. rural).

A protective nesting association with native species counteracts biotic resistance for the spread of an invasive parakeet from urban into rural habitats.

BACKGROUND: Non-native species are often introduced in cities, where they take advantage of microclimatic conditions, resources provided by humans, and competitor/predator release to establish and proliferate. However, native communities in the surrounding rural or natural areas usually halt their spread through biotic resistance, mainly via top-down regulative processes (predation pressure). Here, we show an unusual commensal interaction between exotic and native bird species that favours the spread of the former from urban to rural habitats. RESULTS: We show how Monk parakeets Myiopsitta monachus, an invasive species often introduced in cities worldwide, associated for breeding with a much larger, native species (the white stork Ciconia ciconia) to reduce predation risk in central Spain, thus allowing their colonization of rural areas. Parakeets selected stork nests close to conspecifics and where breeding raptors were less abundant. Parakeets always flushed when raptors approached their nests when breeding alone, but stayed at their nests when breeding in association with storks. Moreover, when storks abandoned a nest, parakeets abandoned it in the following year, suggesting that storks actually confer protection against predators. CONCLUSIONS: Our results show how a protective-nesting association between invasive and native species can counteract biotic resistance to allow the spread of an invasive species across non-urban habitats, where they may become crop pests. Monk parakeet populations are now growing exponentially in several cities in several Mediterranean countries, where they coexist with white storks. Therefore, management plans should consider this risk of spread into rural areas and favour native predators as potential biological controllers.

Isolation and characterization of 15 new microsatellite markers for the globally endangered Lear's macaw Anodorhynchus leari.

A set of 16 microsatellite markers was characterized for Lear's macaw (Anodorhynchus leari) using DNA samples from captive individuals. Extending this molecular toolkit, including the use of samples from wild individuals, is expected to provide the required power of resolution for pedigree inference of both wild and captive individuals, and could support research on the genetic structure of wild populations. We characterize a set of 15 microsatellite markers optimized for the Lear's macaw, developed from a microsatellite-enriched library in a three-step procedure. Primer pairs were initially designed for 62 microsatellites with > 7 tandem repetitions. After amplification of DNA of five wild individuals from different localities, 22 loci seemed to be polymorphic and were further tested on 12 wild nestling samples. Fifteen unlinked loci showed unambiguous peaks and low to moderate polymorphism levels. The combination of the four most polymorphic markers allowed individual identification even of putative sibs.These markers complement previously described microsatellites developed for A. leari and constitute a fundamental genetic toolkit for the investigation of the genetics of both wild and captive populations, thus assisting integrated management plans for the conservation of this globally endangered species.

Climate matching drives spread rate but not establishment success in recent unintentional bird introductions.

Understanding factors driving successful invasions is one of the cornerstones of invasion biology. Bird invasions have been frequently used as study models, and the foundation of current knowledge largely relies on species purposefully introduced during the 19th and early 20th centuries in countries colonized by Europeans. However, the profile of exotic bird species has changed radically in the last decades, as birds are now mostly introduced into the invasion process through unplanned releases from the worldwide pet and avicultural trade. Here we assessed the role of the three main drivers of invasion success (i.e., event-, species-, and location-level factors) on the establishment and spatial spread of exotic birds using an unprecedented dataset recorded throughout the last 100 y in the Iberian Peninsula. Our multimodel inference phylogenetic approach showed that the barriers that need to be overcome by a species to successfully establish or spread are not the same. Whereas establishment is largely related to event-level factors, apparently stochastic features of the introduction (time since first introduction and propagule pressure) and to the origin of introduced species (wild-caught species show higher invasiveness than captive-bred ones), the spread across the invaded region seems to be determined by the extent to which climatic conditions in the new region resemble those of the species' native range. Overall, these results contrast with what we learned from successful deliberate introductions and highlight that different management interventions should apply at different invasion stages, the most efficient strategies being related to event-level factors.

Evolution of genomic variation in the burrowing owl in response to recent colonization of urban areas.

When a species successfully colonizes an urban habitat it can be expected that its population rapidly adapts to the new environment but also experiences demographic perturbations. It is, therefore, essential to gain an understanding of the population structure and the demographic history of the urban and neighbouring rural populations before studying adaptation at the genome level. Here, we investigate populations of the burrowing owl (Athene cunicularia), a species that colonized South American cities just a few decades ago. We assembled a high-quality genome of the burrowing owl and re-sequenced 137 owls from three urban-rural population pairs at 17-fold median sequencing coverage per individual. Our data indicate that each city was independently colonized by a limited number of founders and that restricted gene flow occurred between neighbouring urban and rural populations, but not between urban populations of different cities. Using long-range linkage disequilibrium statistics in an approximate Bayesian computation approach, we estimated consistently lower population sizes in the recent past for the urban populations in comparison to the rural ones. The current urban populations all show reduced standing variation in rare single nucleotide polymorphisms (SNPs), but with different subsets of rare SNPs in different cities. This lowers the potential for local adaptation based on rare variants and makes it harder to detect consistent signals of selection in the genome.

Corticosterone implants produce stress-hyporesponsive birds.

In birds, the use of corticosterone (Cort) implants is a frequent tool aimed at simulating systemic elevations of this hormone and studying effects on biological traits (e.g. physiology, morphology, behavior). This manipulation may alter adrenocortical function, potentially changing both baseline (CortBAS) and stress-induced (CortSTRESS) plasma Cort levels. However, implant effects on the latter trait are rarely measured, disregarding downstream consequences of potentially altered stress responses. Here, we analyzed the effects of Cort implants on both CortBAS and CortSTRESS in nestling and adult European white storks, Ciconia ciconia In addition, we performed a review of 50 studies using Cort implants in birds during the last two decades to contextualize stork results, assess researchers' patterns of use and infer current study biases. High and low doses of Cort implants resulted in a decrease of both CortBAS (31-71% below controls) and CortSTRESS (63-79% below controls) in storks. Our literature review revealed that CortBAS generally increases (72% of experiments) whereas CortSTRESS decreases (78% of experiments) following implant treatment in birds. Our results challenge and expand the prevailing assumption that Cort implants increase circulating CortBAS levels because: (i) CortBAS levels show a quadratic association with implant dose across bird species, and decreased levels may occur at both high and low implant doses, and (ii) Cort implants also decrease CortSTRESS levels, thus producing stress-hyporesponsive phenotypes. It is time to work towards a better understanding of the effects of Cort implants on adrenocortical function, before addressing downstream links to variation in other biological traits.

Shared genetic diversity across the global invasive range of the monk parakeet suggests a common restricted geographic origin and the possibility of convergent selection.

While genetic diversity is hypothesized to be an important factor explaining invasion success, there is no consensus yet on how variation in source populations or demographic processes affects invasiveness. We used mitochondrial DNA haplotypic and microsatellite genotypic data to investigate levels of genetic variation and reconstruct the history of replicate invasions on three continents in a globally invasive bird, the monk parakeet (Myiopsitta monachus). We evaluated whether genetic diversity at invasive sites could be explained by (i) the native source populations from which they were derived and (ii) demographic bottlenecks during introduction. Genetic data indicated a localized source area for most sampled invasive populations, with limited evidence for admixing of native source populations. This pattern largely coincides with historical data on pet trade exports. However, the invasive populations are genetically more similar than predicted from the export data alone. The extent of bottleneck effects varied among invasive populations. The observed low genetic diversity, evidence of demographic contraction and restricted source area do not support the hypothesis that invasion is favoured by the mixing and recombining of genetic variation from multiple source populations. Instead, they suggest that reduced genetic variation through random processes may not inhibit successful establishment and invasion in this species. However, convergent selection across invasive sites could also explain the observed patterns of reduction and similarity in genetic variation and/or the restricted source area. In general, the alternative explanation of intraspecific variation in invasive potential among genotypes or geographic areas is neglected, but warrants more attention as it could inform comparative studies and management of biological invaders.

Can synchronizing feather-based measures of corticosterone and stable isotopes help us better understand habitat-physiology relationships?

Physiological mechanisms link the environment with population dynamics, and glucocorticoid hormones are of particular interest because they respond adaptively to environmental change and can influence vertebrate reproduction and fitness. We tested a novel approach of synchronizing feather-based measures of corticosterone (the primary avian glucocorticoid; CORTf) and ratios of stable isotopes (SIs) of C (δ(13)C) and N (δ(15)N) to provide information about environmental conditions and an integrated physiological response to those conditions over the same period of feather synthesis. Using a fragmented metapopulation of Dupont's larks Chersophilus duponti, an endangered steppe songbird, we analyzed interrelationships among CORTf, δ(13)C, δ(15)N, and the physical environment, including measures of habitat loss and fragmentation. CORTf was not related to any habitat variable measured directly. However, we detected a significant spatial structure to CORTf values and food availability, with greater similarity in both at smaller spatial scales. Using SIs as proxies for the local environment, we found CORTf was negatively related to δ(13)C. Values of CORTf, δ(13)C, and the relationship between the two were likely driven by variation in agricultural land use surrounding lark habitat patches. Our feather-based approach revealed that individual physiology was sensitive to environmental conditions (e.g., an interaction of food availability and variation in habitat) at a local scale, but not patch or landscape scales. Combining CORTf and SIs may be a promising tool because it can provide individual-based information about habitat, physiology, and their relationship during the same time period.

Correction: Sex, personality and conspecific density influence natal dispersal with lifetime fitness consequences in urban and rural burrowing owls.

[This corrects the article DOI: 10.1371/journal.pone.0226089.].

Intraspecific competition and individual behaviour but not urbanization affect the dietary patterns of a generalist avian predator.

Urbanization has reshaped ecosystems and changed natural processes, driving an intense transformation of biomes, biotic community composition and diversity. Despite the growing interest in studying urban ecology over the last decades, the consequences of these changes on species occupying these ecosystems are not yet fully understood. Trophic generalism and tolerance to human disturbance have been proposed as two key traits in the colonization of urban environments. However, most studies focused on species' average traits, paying less attention to the potential role of inter-individual variability. Here, we examined diet specialization in urban and rural breeding pairs, as well as its relationship with individual behaviour and intraspecific competition, using the burrowing owl as a study model. Our results show that both urban and rural breeding pairs behaved as trophic specialists. The diet of burrowing owl breeding pairs followed a gradient from coleopteran- to micromammal-dominated, which is related to individual behaviour (bolder individuals consuming more coleopterans than shyer ones). Besides, pairs distant from others showed a more diverse diet than those experiencing higher levels of intraspecific competition. Models fitted separately for each habitat showed that the proportion of micromammals in the diet of urban breeding pairs was related to their behavior, while the diet of rural pairs was not affected by individual behavior but by intraspecific competition. However, despite the strong selection of tame and more explorative individuals in urban environments and the higher density they reach in this habitat type, they did not differ in their degree of diet specialization from rural conspecifics. Although it would be necessary to evaluate prey availability on a small scale, our results suggest that burrowing owl breeding pairs behave as specialists, despite the generalist character of the species, and that this specialization is not affected by the occupation of urban environments but to individual behaviour and intraspecific competition.

Nesting innovations in neotropical parrots associated to anthropogenic environmental changes.

Parrots are among the most diverse and widely distributed groups of birds and one of the most threatened bird orders mainly due to habitat loss and illegal poaching. Most parrots are obligate cavity nesters, so the logging of mature trees and the transformation of natural cliffs represent important threats to their conservation. Here, we report novel observations of Neotropical parrots nesting in previously unrecorded substrates. We show the first documented case of the cliff-nesting burrowing parrots trying to breed at ground level in an abandoned burrowing owl cavity. Additionally, we provide the first documented observations of this species attempting to nest in building cavities in three urbanized areas of Argentina. Moreover, we report data from four countries of 148 pairs of eight species typically breeding in tree cavity using palm tree bracts as nest sites. Behavioral plasticity in nest sites may allow parrots to maximize their nesting success by exploiting alternative breeding substrates. Moreover, these novelties could contribute to cope with habitat loss and further transformation. However, further research is needed to assess the consequences of these nesting innovations in terms of individual fitness and population dynamics as well as potential factors promoting their appearance.

Lifetime fitness correlates of natal dispersal distance in a colonial bird.

1. Obtaining empirical evidence of the consequences of dispersal distance on fitness is challenging in wild animals because long-term, unbiased data on reproduction, survival and movement are notoriously difficult to obtain. 2. Lifetime fitness correlates of natal dispersal distance were studied in an isolated population of the facultatively colonial lesser kestrel Falco naumanni (Fleischer) monitored during 8 years at north-eastern Spain, where most birds (83%) dispersed from their natal colony to settle at distances ranging from 112 m to 136.5 km. 3. Neither annual breeding success nor age at recruitment was affected by natal dispersal distance. However, a capture-mark-recapture analysis revealed that survival during the year following recruitment decreased exponentially with dispersal distance, with differences of up to 15% between philopatrics and long-distance dispersers. In subsequent years, it remained similar irrespective of the natal dispersal distance moved. These results did not seem to be biased by long-distance dispersers settling differentially in the periphery of the population (which could emigrate permanently and be considered dead in future occasions) or within-individual consistency in successive dispersal distances, so our results appear to reflect genuine survival differences between dispersal tactics. 4. Average lifetime fledgling production, average lifetime recruitment success and rate-sensitive individual fitness (λ(ind)) also decreased with the distance from the natal to the first-breeding colony, indicating that dispersal decisions early in life affecting immediate survival prospects may translate into long-term fitness costs. 5. Both survival and lifetime fitness models including continuous dispersal distances significantly improved the characterization of the effect on fitness compared with models considering dispersal as a discrete process (i.e. dispersal vs. philopatry at a colony level). 6. Long-distance dispersers were more likely to establish new colonies regardless of whether they recruited in the centre or the periphery of the population, revealing their important role in the colonization of unoccupied patches. Individuals experienced a higher probability of mortality in small and newly funded colonies, so lifetime fitness costs of dispersal seem to be explained by recruitment in sites where average quality is low because of high uncertainty in survival prospects.

Apparent Lack of Circovirus Transmission from Invasive Parakeets to Native Birds.

The transmission of pathogens to native species has been highlighted as one of the most important impacts of biological invasions. In this study, we evaluated the presence of psittacine beak and feather disease virus (BFDV) and other circoviruses in native bird species cohabiting with invasive populations of wild rose-ringed (Psittacula krameri) and monk parakeets (Myiopsitta monachus) that were found positive for a particular BFDV genotype in Sevilla, southern Spain. None of the 290 individuals from the 18 native bird species captured showed typical signs of disease caused by BFDV. A sample of 79 individuals from 15 native species showed negative results for the presence of the BFDV genotype previously detected in the sympatric invasive parakeets, as well as any other of the circoviruses tested. Although preliminary, this study suggests a lack of circovirus transmission from invasive parakeets to native birds at the study site. Further research is needed to determine if this apparent absence in transmission depends on the BFDV genotype present in the parakeets, which requires additional screening in other invasive and native populations living in sympatry.

Annual Censuses and Citizen Science Data Show Rapid Population Increases and Range Expansion of Invasive Rose-Ringed and Monk Parakeets in Seville, Spain.

Population changes of invasive species can go unnoticed long before population explosions, so long-term monitoring programs are needed to assess changes in population size. Although invasive populations of rose-ringed (Psittacula krameri) and monk parakeets (Myiopsitta monachus) are present worldwide, their current status and dynamics are mostly poorly known. Here, we provide a long-term population monitoring of both parakeet species established in a Mediterranean urban area. Between 2013 and 2021, we conducted systematic population censuses in the city of Seville and collected their occurrence and spatial distribution data from citizen science platforms. Our censuses showed a rapid population growth of both species: rose-ringed parakeets increased from 1200 to 6300 individuals, while monk parakeets increased from 70 to 1487 individuals. These population trends were weakly reflected by the number of parakeet observations and the number of cells with parakeet observations but not by the number of individuals recorded in citizen science platforms. Moreover, for the monk parakeet, the number of cells with observations was related to the spatial spread of its nests across the study area. Although resource-intensive, long-term monitoring programs are essential to assess population changes and develop effective management actions for invasive species. Thus, contrasting this information with data taken through citizen science platforms can validate the utility of the latter for assessing population status of invasive species.

Nesting innovations allow population growth in an invasive population of rose-ringed parakeets.

Certain traits of recipient environments, such as the availability of limiting resources, strongly determine the establishment success and spread of non-native species. These limitations may be overcome through behavioral plasticity, allowing them to exploit alternative resources. Here, we show how a secondary cavity nester bird, the rose-ringed parakeet Psittacula krameri, innovates its nesting behavior as a response to the shortage of tree cavities for nesting in its invasive range in Tenerife (Canary Islands). We observed that some breeding pairs excavated their own nest cavities in palms, thus becoming primary cavity nester, whereas others occupied nests built with wood sticks by another invasive species, the monk parakeet Myiopsitta monachus. The use of these novel nesting strategies increased the number of breeding pairs by up to 52% over 6 years, contributing to a 128.8% increase of the whole population. Innovative nests were located at greater heights above ground and were more aggregated around conspecifics but did not result in greater breeding success than natural cavities. Occupation of monk parakeet colonies by rose-ringed parakeets also benefited the former species through a protective-nesting association against nest predators. Our results show how an invasive species innovate nesting behaviors and increase nest-site availability in the recipient environment, thus facilitating its population growth and invasion process. Potential behavioral innovations in other invasive rose-ringed parakeet populations may be overlooked, and should be considered for effective management plans.