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Estimating abundance and population size is essential for many ecological and conservation studies of parrots. Achieving these goals requires methods that yield reliable estimates, but parrot traits can make them difficult to detect, count, and capture. We review established and emergent sampling and analytical methods used to estimate parrot abundance and population size, focusing on their assumptions, requirements, and limitations. Roost surveys are cost-effective if all roost locations in a region are known and stable, which is uncommon. Capture-recapture methods incorporate detection probability, but capturing, marking and resighting parrots can be difficult. Distance sampling estimates detection probability and surveys multiple species simultaneously, but is sensitive to the spatial distribution of individuals and excludes birds in flight. Roadside transects can cover large areas and survey multiple species, but habitats near roads may differ from the surrounding areas, biasing abundance estimates. Occupancy surveys and hierarchical models usually require spatially and temporally replicated datasets. Both allow estimation of detection probability; the former dispenses with count data, while the latter is a versatile set of methods that can incorporate multiple processes influencing detection and abundance. Finally, passive acoustic surveys can sample multiple species simultaneously, but identification of vocalisations can be difficult and time-consuming.

With the exception of a few groups of birds, such as large raptors and colonial seabirds, direct counts of nests cannot be conducted over very large areas for most of the abundant and widely distributed species, and thus indirect methods are used to estimate their relative abundances and population sizes. However, many species of the Family Hirundinidae (swallows and martins) build their mud nests in discrete, predictable and accessible sites, which are reused across years. Therefore, the direct count of active nests could constitute a reliable method for estimating breeding population sizes and their changes at large spatial and temporal scales. We illustrate the feasibility of this monitoring approach through a single year survey of >2700 nests of three coexisting Old-World species, the barn swallow ( Hirundo rustica ), the red-rumped swallow ( Cecropis daurica ), and the crag martin ( Ptyonoprogne rupestris ), distributed across Portugal and Spain. Our results revealed changes in the use of nesting substrates and increases in interspecific nest usurpation rates over recent decades. While 56% of the nests of C. daurica were located in rocks five decades ago, almost 100% are nowadays located in anthropogenic substrates such as bridges, road culverts, and abandoned buildings, which could have favored the range expansion of this species. Nest occupation rates were surprisingly low (12% in C. daurica , 21% in H. rustica , and 37% in P. rupestris ), and the proportion of abandoned nesting sites was very high (65% in C. daurica , 50% in H. rustica , and 27% in P. rupestris ). Abandonment rates reflect the population decline reported for H. rustica . Notably, the usurpation of nests of C. daurica by house sparrows Passer domesticus, which is the main cause of breeding failure, has increased from 2.4% in 1976-1979 to 34.7% of the nests nowadays. The long-term monitoring of nests may constitute a reliable and affordable method, with the help of citizen science, for assessing changes in breeding population sizes and conservation threats of these and other mud-nest building hirundines worldwide., Competing Interests: The authors declare that there are no conflicts of interest., (© 2024 The Author(s). Ecology and Evolution published by John Wiley & Sons Ltd.)

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., (© 2023 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

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., Competing Interests: The authors declare no conflict of interest., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Variation in offspring sex ratio, particularly in birds, has been frequently studied over the last century, although seldom using long-term monitoring data. In raptors, the cost of raising males and females is not equal, and several variables have been found to have significant effects on sex ratio, including food availability, parental age, and hatching order. Sex ratio differences between island populations and their mainland counterparts have been poorly documented, despite broad scientific literature on the island syndrome reporting substantial differences in population demography and ecology. Here, we assessed individual and environmental factors potentially affecting the secondary sex ratio of the long-lived Egyptian vulture Neophron percnopterus . We used data collected from Spanish mainland and island populations over a ca. 30-year period (1995-2021) to assess the effects of insularity, parental age, breeding phenology, brood size, hatching order, type of breeding unit (pairs vs. trios), and spatial and temporal variability on offspring sex ratio. No sex bias was found at the population level, but two opposite trends were observed between mainland and island populations consistent with the island syndrome. Offspring sex ratio was nonsignificantly female-biased in mainland Spain (0.47, n  = 1112) but significantly male-biased in the Canary Islands (0.55, n  = 499), where a male-biased mortality among immatures could be compensating for offspring biases and maintaining a paired adult sex ratio. Temporal and spatial variation in food availability might also have some influence on sex ratio, although the difficulties in quantifying them preclude us from determining the magnitude of such influence. This study shows that insularity influences the offspring sex ratio of the Egyptian vulture through several processes that can affect island and mainland populations differentially. Our research contributes to improving our understanding of sex allocation theory by investigating whether sex ratio deviations from parity are possible as a response to changing environments comprised by multiple and complexly interrelated factors., Competing Interests: The authors declare no conflict of interest., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

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., (© 2023. The Author(s).)

In fragile ecosystems, the introduction of exotic species could alter some ecological processes. The Austral parakeet ( Enicognathus ferrugineous ) shows close ecological and evolutionary relationships with the Andean Araucaria ( Araucaria araucana ), so any alteration in these interactions may have negative consequences for both partners and for ecosystem functioning and structure. We conducted extensive roadside surveys to estimate the abundance of parakeets in the northern Patagonian Andes over 4 years and recorded the food plants consumed by foraging flocks. The use of native habitats and humanized areas like villages and farms was influenced by the Araucaria seed crop. In masting years, the large seed crop allowed a massive use of this resource during the non-breeding season, and even during the breeding season. The exploitation of exotic plants was minor in the masting year, but became predominant in non-masting years, especially during the non-breeding season. This feeding switch towards exotic plants primarily arose because the low Araucaria seed crop in non-masting years is entirely consumed just after production by domestic and wild exotic mammals living in Araucaria forests year-round, thus forcing the displacement of parakeets towards anthropic habitats to exploit exotic plants. Given the degradation of the remaining Andean Araucaria forests due to the impact of exotic mammals on the ecological interaction between Araucaria and Austral parakeets, ambitious programs to exclude or reduce the density of these alien mammals, including livestock, are warranted., Competing Interests: All authors declare no competing interests., (© 2022 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Urbanization creates new ecological conditions that can affect biodiversity at all levels, including the diversity and prevalence of parasites of species that may occupy these environments. However, few studies have compared bird-ectoparasite interactions between urban and rural individuals. Here, we analyze the ectoparasite community and co-infection patterns of urban and rural burrowing owls, Athene cunicularia , to assess the influence of host traits (i.e., sex, age, and weight), and environmental factors (i.e., number of conspecifics per nest, habitat type and aridity) on its composition. Ectoparasites of burrowing owls included two lice, one flea, and one mite. The overall prevalence for mites, lice and fleas was 1.75%, 8.76% and 3.50%, respectively. A clear pattern of co-infection was detected between mites and fleas and, to less extent, between mites and lice. Adult owls harbored fewer ectoparasites than nestlings, and adult females harbored more lice than males. Our results also show that mite and flea numbers were higher when more conspecifics cohabited the same burrow, while lice showed the opposite pattern. Rural individuals showed higher flea parasitism and lower mite parasitism than urban birds. Moreover, mite numbers were negatively correlated with aridity and host weight. Although the ectoparasitic load of burrowing owls appears to be influenced by individual age, sex, number of conspecifics per nest, and habitat characteristics, the pattern of co-infection found among ectoparasites could also be mediated by unexplored factors such as host immune response, which deserves further research.

Wildlife trade is a major driver of biodiversity loss worldwide. To regulate its impact, laws and regulations have been implemented at the international and national scales. The Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) has regulated the international legal trade since 1975. However, an important volume of illegal trade-mainly within countries-continues to threaten several vertebrate groups, which could be due to a lack of specific legislation or enforcement of existing regulations. Our aim was to gain a more accurate picture of poaching and legal possession of native parrots as pets in the Neotropics, where illegal domestic trade is currently widespread. We conducted a systematic search of the laws of each of the 50 countries and overseas territories, taking into account their year of implementation and whether the capture, possession and/or sale of parrots is permitted. We compared this information with legal exports reported by CITES to assess differences between the enforcement of international and national trade regulations. We found that only two countries (Guyana and Suriname) currently allow the capture, trade and possession of native parrots, while Peru allowed international legal trade until recently. The other countries have banned parrot trade from years to decades ago. However, the timing of implementation of international and national trade regulations varied greatly between countries, with half of them continuing to export parrots legally years or decades after banning domestic trade. The confusion created by this complex legal system may have hindered the adoption of conservation measures, allowing poaching, keeping and trade of protected species within and between neighboring countries. Most countries legally exported Neotropical parrot species which were not native to those countries, indicating that trans-border smuggling often occurred between neighboring countries prior to their legal exportations, and that this illicit activity continues for the domestic trade. Governments are urged to effectively implement current legislation that prohibits the trapping and domestic trade of native parrots, but also to develop coordinated alliances and efforts to halt illegal trade among them. Otherwise, illegal trade will continue to erode the already threatened populations of a large number of parrot species across the Neotropics.

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.

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.

The global pet trade is a major pathway for the introduction of invasive alien species. The composition of species selected for transport is driven by market demands, which may be influenced by a combination of both historical and cultural factors. We compared Eastern (Taiwan) and Western (Australia and the Iberian Peninsula) bird markets to explore factors associated with the species composition and geographic origin of the birds for sale. We used a bespoke randomization test to compare species composition, geographic origins, and species overlap at different taxonomic levels among bird markets across countries. Alien species identified in the study accounted for more than 10% of the world's bird species. Parrots and songbirds were the most common alien bird taxa traded across all markets. In both Iberian and Australian markets, there was a strong bias toward parrots, waxbills, gamebirds, and finches. In Taiwan, species traded more than expected were parrots, waxbills, starlings, and leafbirds. Neotropical species were the most traded group in the three markets. Afrotropical species were also traded more than expected in Iberian and Australian markets, while the Taiwanese traded more alien species from neighboring Asian regions. The bird trade focuses on the same few bird groups worldwide. The composition and origin of species preferred in the Western markets may be influenced by colonial histories, cultural similarity, and strict regulations on wildlife importation, while species preferences in Eastern markets are strongly influenced by regional culture and proximity. Propagule pressure is a dominant factor influencing the success of biological invasions; it is important to recognize differences in the composition of bird markets among regions because they can translate into different invasion risks, among other factors., Competing Interests: None of the authors have any competing interests., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

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., (© The Author(s) (2021). Published by Oxford University Press on behalf of Editorial Office, Current Zoology.)

While most of the knowledge on invasive species focuses on their impacts, little is known about their potential positive effects on other species. Invasive ecosystem engineers can disrupt recipient environments; however, they may also facilitate access to novel resources for native species. The monk parakeet ( Myiopsitta monachus ) is a worldwide invader and the only parrot that builds its own communal nests, which can be used by other species. However, the ecological effects of these interspecific interactions are barely known. We compared the role of the monk parakeet as a nest-site facilitator in different rural and urban areas, both invaded and native, across three continents and eight breeding seasons. A total of 2690 nests from 42 tenant species, mostly cavity-nesting birds, were recorded in 26% of 2595 monk parakeet nests. Rural and invaded areas showed the highest abundance and richness of tenant species. Multispecies communal nests triggered interspecific aggression between the monk parakeet host and its tenants, but also a cooperative defense against predators. Despite the positive effects for native species, monk parakeets also facilitate nesting opportunities to other non-native species and may also transmit diseases to tenants, highlighting the complexity of biotic interactions in biological invasions.

Plant-animal interactions are key to sustaining whole communities and ecosystem function. However, their complexity may limit our understanding of the underlying mechanisms and the species involved. The ecological effects of epizoochory remain little known compared to other seed dispersal mechanisms given the few vectors identified. In addition, epizoochory is mostly considered non-mutualistic since dispersers do not obtain nutritional rewards. Here, we show a widespread but unknown mutualistic interaction between parrots and plants through epizoochory. Combining our observations with photos from web-sources, we recorded nearly 2000 epizoochory events in 48 countries across five continents, involving 116 parrot species and nearly 100 plant species from 35 families, including both native and non-native species. The viscid pulp of fleshy fruits and anemochorous structures facilitate the adherence of tiny seeds (mean 3.7 × 2.56 mm) on the surface of parrots while feeding, allowing the dispersion of these seeds over long distances (mean = 118.5 m). This parrot-plant mutualism could be important in ecosystem functioning across a wide diversity of environments, also facilitating the spread of exotic plants. Future studies should include parrots for a better understanding of plant dispersal processes and for developing effective conservation actions against habitat loss and biological invasions.

Natal dispersal, the movement between the birth and the first breeding site, has been rarely studied in long-lived territorial birds with a long-lasting pre-breeding stage. Here we benefited from the long-term monitoring programs of six populations of Egyptian vultures (Neophron percnopterus) from Spain and France to study how the rearing environment determines dispersal. For 124 vultures, we recorded a median dispersal distance of 48 km (range 0-656 km). Linear models were used to assess the effect of population and individual traits on dispersal distance at two spatial scales. Dispersal distances were inversely related to vulture density in the natal population, suggesting that birds perceive the abundance of conspecifics as a signal of habitat quality. This was particularly true for declining populations, so increasing levels of opportunistic philopatry seemed to arise in high density contexts as a consequence of vacancies created by human-induced adult mortality. Females dispersed further than males, but males were more sensitive to the social environment, indicating different dispersal tactics. Both sexes were affected by different individual attributes simultaneously and interactively with this social context. These results highlight that complex phenotype-by-environment interactions should be considered for advancing our understanding of dispersal dynamics in long-lived organisms.

Behavioural and socio-cultural traits are recognized in the restriction of gene flow in species with high cognitive capacity and complex societies. This isolation by social barriers has been generally overlooked in threatened species by assuming disrupted gene flow due to population fragmentation and decline. We examine the genetic structure and ecology of the global population of the Critically Endangered red-fronted macaw (Ara rubrogenys), an endemic species to the inter-Andean valleys of Bolivia. We found a fine-scale genetic structuring in four genetic clusters. Genetic diversity was higher in wild compared to captive-bred macaws, but similar to that of captive wild-caught macaws. We found no clear evidence of severe genetic erosion in the population in recent decades, but it was patent in historic times, overlapping with drastic human habitat transformation and macaw persecution over millennia. We found no evidence of geographical and ecological barriers, owing to the high dispersal ability, nesting and foraging habits between genetic clusters. The lack of genetic intermixing despite long-distance foraging and seasonal movements suggests recruitment in natal colonies and other social factors reinforcing philopatry-related genetic structure. Conservation efforts should be specifically focussed on major threats in each genetic cluster as independent conservation units, and also considered in ex-situ management.

In some vertebrate species, family units are typically formed when sexually mature individuals delay dispersal and independent breeding to remain as subordinates in a breeding group. This behaviour has been intensively studied in gregarious species but has also been described in non-social species where ecological and evolutionary drivers are less known. Here, we explore factors that favour delayed dispersal and family living and potential benefits associated with this strategy in a non-social, monogamous species (the burrowing owl, Athene cunicularia) occupying urban and rural habitats. Our results show that family units arise when first-year individuals, mainly males, delay their dispersal to stay in their natal nests with their parents. This delayed dispersal, while still uncommon, was more prevalent in urban (7%) than in rural (3%) habitats, and in areas with high conspecific density and productivity. Birds delaying dispersal contributed to the genetic pool of the offspring in 25% of the families analysed, but did not increase the productivity of the nests where they remained. However, their presence was related to an improvement in the body condition of chicks, which was ultimately linked to a slightly positive effect in offspring future survival probabilities. Finally, delayed dispersers were recruited as breeders in high-quality urban territories and closer to their natal nests than individuals dispersing during their first year of life. Thus, our results suggest that delaying dispersal may be mainly related to opportunities to inheriting a good quality territory, especially for males. Our study contributes to understanding the role played by habitat quality in promoting delayed dispersal and family living, not only in social but also non-social species, highlighting its impact in the ecology and evolution of animal populations.

Biological invasion is a global problem with large negative impacts on ecosystems and human societies. When a species is introduced, individuals will first have to pass through the invasion stages of uptake and transport, before actual introduction in a non-native range. Selection is predicted to act during these earliest stages of biological invasion, potentially influencing the invasiveness and/or impact of introduced populations. Despite this potential impact of pre-introduction selection, empirical tests are virtually lacking. To test the hypothesis of pre-introduction selection, we followed the fate of individuals during capture, initial acclimation, and captivity in two bird species with several invasive populations originating from the international trade in wild-caught pets (the weavers Ploceus melanocephalus and Euplectes afer ). We confirm that pre-introduction selection acts on a wide range of physiological, morphological, behavioral, and demographic traits (incl. sex, age, size of body/brain/bill, bill shape, body mass, corticosterone levels, and escape behavior); these are all traits which likely affect invasion success. Our study thus comprehensively demonstrates the existence of hitherto ignored selection acting before the actual introduction into non-native ranges. This could ultimately change the composition and functioning of introduced populations, and therefore warrants greater attention. More knowledge on pre-introduction selection also might provide novel targets for the management of invasive species, if pre-introduction filters can be adjusted to change the quality and/or quantity of individuals passing through such that invasion probability and/or impacts are reduced., Competing Interests: None declared., (© 2020 The Authors. Evolutionary Applications published by John Wiley & Sons Ltd.)

Background: Invasive Africanized honey bees potentially compete with cavity-nesting birds in South America. However, the impacts of this competition and its conservation consequences to threatened species are poorly known. We quantified the presence of these bees and assessed their competition for cliff cavities used by nesting Lear's macaws Anodorhynchus leari, a globally endangered parrot endemic to the Caatinga biome of Brazil. We treated beehives with permethrin by shooting them with a crossbow bolt that distributed the compound upon impact. When feasible, we removed the comb and applied an insecticide (fipronil) to deter bee recolonization. We subsequently surveyed the macaw breeding population to verify whether our treatment allowed for nest recruitment in cavities previously occupied by bees., Results: We recorded > 100 beehives in the nesting cliffs. Hives outnumbered macaw nests tenfold in two areas recently recolonized by macaws. Cavities occupied by bees were significantly higher than those occupied by macaws, suggesting that macaws may be forced to breed in lower cavities. None of the untreated cavities (n = 50) were occupied by nesting macaws, whereas 15% of treated cavities (n = 52) were occupied within 2 years post treatment. Treated cavities occupied by macaws were significantly higher than those not occupied. Hive management increased macaw breeding population by 71% of the macaw breeding population increase., Conclusion: Experimental hive treatments were effective in restoring nesting resources lost due to bee infestation. An intensive and continued eradication program is recommended to enhance macaw habitat restoration, facilitating its expansion into historical areas. © 2020 Society of Chemical Industry., (© 2020 Society of Chemical Industry.)

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.

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)., (© 2020 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

The hypothalamus-pituitary-adrenal/interrenal (HPA) axis of vertebrates integrates external information and orchestrates responses to cope with energy-demanding and stressful events through changes in circulating glucocorticoid levels. Urbanization exposes animals to a wide variety of ever-changing stimuli caused by human activities that may affect local wildlife populations. Here, we empirically tested the hypothesis that urban and rural owls ( Athene cunicularia ) show different adrenocortical responses to stress, with urban individuals showing a reduced HPA-axis response compared to rural counterparts to cope with the high levels of human disturbance typical of urban areas. We applied a standard capture-restraint protocol to measure baseline levels and stress-induced corticosterone (CORT) responses. Urban and rural owls showed similar circulating baseline CORT levels. However, maximum CORT levels were attained earlier and were of lower magnitude in urban compared to rural owls, which showed a more pronounced and long-lasting response. Variability in CORT responses was also greater in rural owls and contained the narrower variability displayed by urban ones. These results suggest that only individuals expressing low-HPA-axis responses can thrive in cities, a pattern potentially mediated by three alternative and non-exclusive hypotheses: phenotypic plasticity, natural selection and matching habitat choice. Due to their different conservation implications, we recommend further research to properly understand wildlife responses to humans in an increasingly urbanized world., (© The Author(s) 2020. Published by Oxford University Press and the Society for Experimental Biology.)

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., Competing Interests: Competing interestsThe authors declare that they have no competing interests., (© The Author(s) 2020.)

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.

There is a growing need to understand how species respond to habitat changes and the potential key role played by natal dispersal in population dynamics, structure and gene flow. However, few studies have explored differences in this process between conspecifics living in natural habitats and those inhabiting landscapes highly transformed by humans, such as cities. Here, we investigate how individual traits and social characteristics can influence the natal dispersal decisions of burrowing owls (Athene cunicularia) living in urban and rural areas, as well as the consequences in terms of reproductive success and apparent survival. We found short dispersal movements among individuals, with differences between urban and rural birds (i.e., the former covering shorter distances than the latter), maybe because of the higher conspecific density of urban compared to rural areas. Moreover, we found that urban and rural females as well as bold individuals (i.e., individuals with shorter flight initiation distance) exhibited longer dispersal distances than their counterparts. These dispersal decisions have effects on individual fitness. Individuals traveling longer distances increased their reproductive prospects (productivity during the first breeding attempt, and long term productivity). However, the apparent survival of females decreased when they dispersed farther from their natal territory. Although further research is needed to properly understand the ecological and evolutionary consequences of dispersal patterns in transformed habitats, our results provide information about the drivers and the consequences of the restricted natal movements of this species, which may explain its population structuring through restricted gene flow between and within urban and rural areas., Competing Interests: Authors declare that they have neither competing financial and/or non-financial interests nor any confidentiality agreements

Anecdotic citations of food wasting have been described for parrots, but we lack a comprehensive knowledge about the extent of this behaviour, and its ecological and evolutionary implications. Here, we combine experimental and observational approaches to evaluate the spatial, temporal, typological and taxonomic extent of food wasting by parrots, to identify the ecological and evolutionary factors driving food wasting, and to assess the incidence of two ecological functions derived from food wasting, such as food facilitation to other animal species and secondary seed dispersal. We found that food wasting is a widespread behaviour found in all the studied parrot species. However, the proportion of food wasted differed among species and throughout the year. Parrots wasted more food during the non-breeding season, when they relied on exotic plants and on unripe fruits or seeds. We also recorded 86 animal species feeding on the food wasted by parrots, 27 of which potentially acted as secondary seed dispersers. Overall, our study emphasizes the universality of food wasting among parrots, and the important implications that this behaviour may have for the species involved (i.e., the parrot, the plant, the other species feeding on wasted food), and for the functioning of the whole ecosystem.

Although the limited resources available to save species from extinction necessitate the optimization of conservation actions, little is known about their costs and effectiveness. We developed a costs-rewards framework that integrates information on which sectors of society contribute to funding conservation, how much is contributed, how funds are distributed among conservation targets and how these investments drive not only conservation rewards but also the economic and ecosystem services that benefit society. We applied this framework to the Lear's macaw ( Anodorhynchus leari ), a species discovered in the wild in 1978 with only 60 individuals. Funds invested over the last 25 years reached US$3.66 million. The contribution of governments, non-governmental organizations and private funders varied over time, as did the funding targets. Funds were proportionally invested to mitigate the main causes of mortality, while no funds were devoted to protecting foraging habitats. Conservation rewards were satisfactory, with the cost and time needed to downlist the species from critically endangered to endangered being similar to those invested in other bird species. However, economic rewards (through ecotourism and handicrafts linked to the conservation of the species) were low and require promotion, while ecosystem services provided by Lear's macaws have yet to be quantified., Competing Interests: The authors declare that they have no competing interests.

Dispersal propensity has been correlated with personality traits, conspecific density and predation risk in a variety of species. Thus, changes in the relative frequency of behavioural phenotypes or in the ecological pressures faced by individuals in contrasting habitats can have unexpected effects on their dispersal strategies. Here, using the burrowing owl Athene cunicularia as a study model, we test whether changes in the behavioural profile of individuals and changes in conspecific density and predation pressure associated with urban life influence their breeding dispersal decisions compared to rural conspecifics. Our results show that breeding dispersal behaviour differs between rural and urban individuals. Site fidelity was lower among rural than among urban birds, and primarily related to an individual's behaviours (fear of humans), which has been reported to reflect individual personality. In contrast, the main determinant of site fidelity among urban owls was conspecific density. After taking the decision of dispersing, urban owls moved shorter distances than rural ones, with females dispersing farther than males. Our results support a personality-dependent dispersal pattern that might vary with predation risk. However, as multiple individuals of two populations (one urban, one rural) were used for this research, differences can thus also be caused by other factors differing between the two populations. Further research is needed to properly understand the ecological and evolutionary consequences of changes in dispersal behaviours, especially in terms of population structuring and gene flow between urban and rural populations.

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 (Cort BAS ) and stress-induced (Cort STRESS ) 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 Cort BAS and Cort STRESS 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 Cort BAS (31-71% below controls) and Cort STRESS (63-79% below controls) in storks. Our literature review revealed that Cort BAS generally increases (72% of experiments) whereas Cort STRESS decreases (78% of experiments) following implant treatment in birds. Our results challenge and expand the prevailing assumption that Cort implants increase circulating Cort BAS levels because: (i) Cort BAS 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 Cort STRESS 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., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Background: Urbanization is one of the main causes of biodiversity loss worldwide. Wildlife responses to urbanization, however, are greatly variable and, paradoxically, some threatened species may achieve much larger populations in urban than in natural habitats. Urban conservation hotspots may therefore help some species avoid regional or even global extinctions, but not conserve their often overlooked ecological functions in the wild. We aim to draw attention to this issue using two species of globally threatened parrots occurring in the Dominican Republic: the Hispaniolan amazon ( Amazona ventralis ) and the Hispaniolan parakeet ( Psittacara chloropterus )., Methods: We conducted a large-scale roadside survey in June 2017 across the country to estimate the relative abundance of parrots in natural habitats, rural habitats, and cities. We combined this with informal interviews with local people to collect information on past and current human impacts on parrot populations. We also looked for foraging parrots to assess their potential role as seed dispersers, an ecological function that has been overlooked until very recently., Results: Relative abundances of both parrot species were negligible in rural areas and very low in natural habitats. They were generally between one and two orders of magnitude lower than that of congeneric species inhabiting other Neotropical ecosystems. Relative abundances were six times higher in cities than in natural habitats in the case of the Hispaniolan parakeet and three times higher in the case of the Hispaniolan amazon. People indicated hunting for a source food and to mitigate crop damage as causes of parrot population declines, and a vigorous illegal trade for parrots (131 individuals recorded, 75% of them poached very recently), mostly obtained from protected areas where the last small wild populations remain. We observed parrots foraging on 19 plant species from 11 families, dispersing the fruits of 14 species by carrying them in their beaks and consuming them in distant perching trees. They discarded undamaged mature seeds, with the potential to germinate, in 99.5% of cases ( n = 306), and minimum dispersal distances ranged from 8 to 155 m (median = 37 m)., Discussion: The loss of ecological functions provided by some species when they disappear from natural habitats and only persist in cities may have long-term, unexpected effects on ecosystems. Our example demonstrates how two cities may soon be the last refuges for two endemic parrots if overharvesting continues, in which case their overlooked role as seed dispersers would be completely lost in nature. The functional extinction of these species could strongly affect vegetation communities in an island environment where seed-dispersal species are naturally scarce. While conservation plans must include urban populations of threatened species, greater efforts are needed to restore their populations in natural habitats to conserve ecological functions., Competing Interests: The authors declare that they have no competing interests.

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., (© 2018 The Authors.)

The identification of effects of invasive species is challenging owing to their multifaceted impacts on native biota. Negative impacts are most often reflected in individual fitness rather than in population dynamics of native species and are less expected in low-biodiversity habitats, such as urban environments. We report the long-term effects of invasive rose-ringed parakeets on the largest known population of a threatened bat species, the greater noctule, located in an urban park. Both species share preferences for the same tree cavities for breeding. While the number of parakeet nests increased by a factor of 20 in 14 years, the number of trees occupied by noctules declined by 81%. Parakeets occupied most cavities previously used by noctules, and spatial analyses showed that noctules tried to avoid cavities close to parakeets. Parakeets were highly aggressive towards noctules, trying to occupy their cavities, often resulting in noctule death. This led to a dramatic population decline, but also an unusual aggregation of the occupied trees, probably disrupting the complex social behaviour of this bat species. These results indicate a strong impact through site displacement and killing of competitors, and highlight the need for long-term research to identify unexpected impacts that would otherwise be overlooked., Competing Interests: The authors have no competing interests.

PCR is a universal tool for the multiplication of specific DNA sequences. For example, PCR-based sex determination is widely used, and a diversity of primer sets is available. However, this protocol requires thermal cycling and electrophoresis, so results are typically obtained in laboratories and several days after sampling. Loop-mediated isothermal amplification (LAMP) is an alternative to PCR that can take molecular ecology outside the laboratory. Although its application has been successfully probed for sex determination in three species of a single avian Family (raptors, Accipitridae), its generality remains untested and suitable primers across taxa are lacking. We designed and tested the first LAMP-based primer set for sex determination across the modern birds (NEO-W) based on a fragment of the gene chromo-helicase-DNA-binding protein located on the female-specific W chromosome. As nucleotide identity is expected to increase among more related taxa, taxonomically targeted primers were also developed for the Order Falconiformes and Families Psittacidae, Ciconiidae, Estrildidae and Icteridae as examples. NEO-W successfully determined sex in a subset of 21 species within 17 Families and 10 Orders and is therefore a candidate primer for all modern birds. Primer sets designed specifically for the selected taxa correctly assigned sex to the evaluated species. A short troubleshooting guide for new LAMP users is provided to identify false negatives and optimize LAMP reactions. This study represents the crucial next step towards the use of LAMP for molecular sex determination in birds and other applications in molecular ecology., (© 2017 John Wiley & Sons Ltd.)

The exponential growth of the human population often causes the overexploitation of resources and disruption of ecological interactions. Here, we propose that the antagonist effect of humans on exploited species might be alleviated with the advent of a second predator species. We focused on the complex interactions between an endangered conifer ( Araucaria araucana ) and two seed exploiters: the Austral parakeet ( Enicognathus ferrugineus ) and human seed collectors. We tested the importance of partial seed consumption by parakeets as an escape from human seed harvesting. Although parakeets frequently ate whole seeds, a substantial proportion of the seeds found under trees were only partially eaten and avoided by human seed collectors. These seeds germinated at a similar proportion but faster than intact seeds under laboratory conditions. Our results revealed an overlooked mutualism between parakeets and an endangered tree. Incomplete seed eating by parakeets, plus selection against these eaten seeds by humans, may enhance regeneration possibilities for this conifer species subject to human seed collection, turning the scale of the antagonism-mutualism continuum to the mutualistic side. In this context, parakeets might be providing an important service in those forests subject to human harvesting by allowing a fraction of seeds to escape human predation., Competing Interests: We have no competing interests.

Human-induced biological invasions are common worldwide and often have negative impacts on wildlife and human societies. Several studies have shown evidence for selection on invaders after introduction to the new range. However, selective processes already acting prior to introduction have been largely neglected. Here, we tested whether such early selection acts on known behaviour-related gene variants in the yellow-crowned bishop (Euplectes afer), a pet-traded African songbird. We tested for nonrandom allele frequency changes after trapping, acclimation and survival in captivity. We also compared the native source population with two independent invasive populations. Allele frequencies of two SNPs in the dopamine receptor D4 (DRD4) gene-known to be linked to behavioural activity in response to novelty in this species-significantly changed over all early invasion stages. They also differed between the African native population and the two invading European populations. The two-locus genotype associated with reduced activity declined consistently, but strongest at the trapping stage. Overall genetic diversity did not substantially decrease, and there is little evidence for new alleles in the introduced populations, indicating that selection at the DRD4 gene predominantly worked on the standing genetic variation already present in the native population. Our study demonstrates selection on a behaviour-related gene during the first stages of a biological invasion. Thus, pre-establishment stages of a biological invasion do not only determine the number of propagules that are introduced (their quantity), but also their phenotypic and genetic characteristics (their quality)., (© 2017 The Authors. Molecular Ecology Published by John Wiley & Sons Ltd.)

Theory predicts that contrasting properties of mutualistic and antagonistic networks differentially promote community resilience to species loss. However, the outcome of most ecological interactions falls within a continuum between mutualism and antagonism, and we ignore the extent to which this interactions' continuum might influence community stability. Using a large data set of interactions, we compared co-extinction cascades that either consider or ignore the mix of beneficial and detrimental actions that parrots exert on plants. When the antagonism-mutualism continuum was considered, a combination of the properties that separately enhance community stability in ecological networks emerged. This combination of properties led to an overall increase of the parrot community robustness to face plant species loss. Our results highlight that the conditional outcomes of interactions can influence the structure of ecological networks, thus affecting our predictions of community stability against eventual changes.

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., Competing Interests: The authors declare no conflict of interest.