The Mitogenome Relationships and Phylogeography of Barn Swallows (Hirundo rustica).

The barn swallow (Hirundo rustica) poses a number of fascinating scientific questions, including the taxonomic status of postulated subspecies. Here, we obtained and assessed the sequence variation of 411 complete mitogenomes, mainly from the European H. r. rustica, but other subspecies as well. In almost every case, we observed subspecies-specific haplogroups, which we employed together with estimated radiation times to postulate a model for the geographical and temporal worldwide spread of the species. The female barn swallow carrying the Hirundo rustica ancestral mitogenome left Africa (or its vicinity) around 280 thousand years ago (kya), and her descendants expanded first into Eurasia and then, at least 51 kya, into the Americas, from where a relatively recent (<20 kya) back migration to Asia took place. The exception to the haplogroup subspecies specificity is represented by the sedentary Levantine H. r. transitiva that extensively shares haplogroup A with the migratory European H. r. rustica and, to a lesser extent, haplogroup B with the Egyptian H. r. savignii. Our data indicate that rustica and transitiva most likely derive from a sedentary Levantine population source that split at the end of the Younger Dryas (YD) (11.7 kya). Since then, however, transitiva received genetic inputs from and admixed with both the closely related rustica and the adjacent savignii. Demographic analyses confirm this species' strong link with climate fluctuations and human activities making it an excellent indicator for monitoring and assessing the impact of current global changes on wildlife.

Evolutionary dynamics in the Anthropocene: Life history and intensity of human contact shape antipredator responses.

Humans profoundly impact landscapes, ecosystems, and animal behavior. In many cases, animals living near humans become tolerant of them and reduce antipredator responses. Yet, we still lack an understanding of the underlying evolutionary dynamics behind these shifts in traits that affect animal survival. Here, we used a phylogenetic meta-analysis to determine how the mean and variability in antipredator responses change as a function of the number of generations spent in contact with humans under 3 different contexts: urbanization, captivity, and domestication. We found that any contact with humans leads to a rapid reduction in mean antipredator responses as expected. Notably, the variance among individuals over time observed a short-term increase followed by a gradual decrease, significant for domesticated animals. This implies that intense human contact immediately releases animals from predation pressure and then imposes strong anthropogenic selection on traits. In addition, our results reveal that the loss of antipredator traits due to urbanization is similar to that of domestication but occurs 3 times more slowly. Furthermore, the rapid disappearance of antipredator traits was associated with 2 main life-history traits: foraging guild and whether the species was solitary or gregarious (i.e., group-living). For domesticated animals, this decrease in antipredator behavior was stronger for herbivores than for omnivores or carnivores and for solitary than for gregarious species. By contrast, the decrease in antipredator traits was stronger for gregarious, urbanized species, although this result is based mostly on birds. Our study offers 2 major insights on evolution in the Anthropocene: (1) changes in traits occur rapidly even under unintentional human "interventions" (i.e., urbanization) and (2) there are similarities between the selection pressures exerted by domestication and by urbanization. In all, such changes could affect animal survival in a predator-rich world, but through understanding evolutionary dynamics, we can better predict when and how exposure to humans modify these fitness-related traits.

EVI and NDVI as proxies for multifaceted avian diversity in urban areas.

Most ecological studies use remote sensing to analyze broad-scale biodiversity patterns, focusing mainly on taxonomic diversity in natural landscapes. One of the most important effects of high levels of urbanization is species loss (i.e., biotic homogenization). Therefore, cost-effective and more efficient methods to monitor biological communities' distribution are essential. This study explores whether the Enhanced Vegetation Index (EVI) and the Normalized Difference Vegetation Index (NDVI) can predict multifaceted avian diversity, urban tolerance, and specialization in urban landscapes. We sampled bird communities among 15 European cities and extracted Landsat 30-meter resolution EVI and NDVI values of the pixels within a 50-m buffer of bird sample points using Google Earth Engine (32-day Landsat 8 Collection Tier 1). Mixed models were used to find the best associations of EVI and NDVI, predicting multiple avian diversity facets: Taxonomic diversity, functional diversity, phylogenetic diversity, specialization levels, and urban tolerance. A total of 113 bird species across 15 cities from 10 different European countries were detected. EVI mean was the best predictor for foraging substrate specialization. NDVI mean was the best predictor for most avian diversity facets: taxonomic diversity, functional richness and evenness, phylogenetic diversity, phylogenetic species variability, community evolutionary distinctiveness, urban tolerance, diet foraging behavior, and habitat richness specialists. Finally, EVI and NDVI standard deviation were not the best predictors for any avian diversity facets studied. Our findings expand previous knowledge about EVI and NDVI as surrogates of avian diversity at a continental scale. Considering the European Commission's proposal for a Nature Restoration Law calling for expanding green urban space areas by 2050, we propose NDVI as a proxy of multiple facets of avian diversity to efficiently monitor bird community responses to land use changes in the cities.

Defensive tolerance to parasitism is correlated with sexual selection in swallows.

Parasite-mediated sexual selection has been the topic of extensive research and enthusiastic debate for more than three decades. Here, we suggest that secondary sexual characters may not only signal parasite resistance but also defensive tolerance. We exemplify this possibility by analysing information on two sexually selected traits, annual reproductive success, and ectoparasitism in a barn swallow Hirundo rustica population followed for more than 30 years. For each individual, we estimated the slope of the association between reproductive success and parasitism as an index of tolerance and subsequently explored the association with the expression of the sexually selected traits. In accordance with expectations of parasites playing a role in sexual selection, tail length was negatively related to load of chewing lice and nest size was positively related to tolerance to chewing lice. We discuss the importance of considering defensive tolerance for understanding the role of parasite-mediated sexual selection.

Long-term consistency in susceptibility of prey species to predation by an avian predator.

Selection by predators affects prey through competition for limiting resources. This not only has consequences for direct mortality but also indirectly affects disturbance. Changes in the intensity of selection on prey by predators may affect the size of prey populations, with consequences for their short- or long-term interactions. We assessed whether predation by northern goshawks Accipiter gentilis modified the composition of prey communities consistently along a temporal gradient, showing long-term consistency in susceptibility of prey species to predation. We followed six populations of the goshawk in two biomes in Denmark and Finland during 1949-2019. Susceptibility to goshawk predation in 2005-2017 in Denmark was only weakly related to susceptibility to goshawk predation in 1977-2004. In Finland, susceptibility of shared prey species to goshawk predation was positively related between periods. The average difference in susceptibility to goshawk predation between periods was considerably higher in Denmark than in Finland. Susceptibility of prey species to predation in goshawks increased with latitude and body mass of prey species, and decreased with period of time and population density of prey species. The changes in susceptibility to predation suggest changes in the characteristics of the local prey pools.

Nest size matters: common cuckoos prefer to parasitize larger nests of Oriental reed warblers.

Avian brood parasites leave parental care of their offspring to foster parents. Theory predicts that parasites should select for large host nests when they have sufficient available host nests at a given time. We developed an empirical experimental design to test cognitive ability of female cuckoos in nest size by studying nest choice of common cuckoos (Cuculus canorus) among nests of its Oriental reed warbler (Acrocephalus orientalis) hosts. We presented three groups of experimental nests: 1) nest dyads tied together including one large and one small artificial nest from reed leaves, 2) nest triads tied together used the old modified warbler's own nests including enlarged, reduced and medium-sized nests, and 3) nest dyads are similar to group 1, but not tied together to elicit parasitism by common cuckoos. We predict that cuckoos prefer larger nest than medium one, the next is smaller nest. Our findings showed that common cuckoo females generally prefer large nests over medium or small sized nests in all three experimental groups. Furthermore, cuckoo parasitism was significantly more common than in previous studies of the same warbler population, implying that larger, higher and more exposed host nests effectively increased the probability of cuckoo parasitism.

Long-term trends in the phylogenetic and functional diversity of Anatidae in South China coastal wetlands.

Species loss has attracted much attention among scientists for more than half a century. However, we have little information on the trends in phylogenetic and functional changes behind the species loss although this information is always asynchronous and important for conservation and management. We measured community trends in Anatidae (ducks and geese) for the last 50 yr to quantify trends in phylogenetic and functional diversity patterns coinciding with taxonomic historical dynamics. We used one-way ANOVAs to test if there was a significant historical trend in communities of Anatidae. We characterized taxonomic, phylogenetic, and functional diversity of communities. For taxonomic diversity, we used species richness (SR). For phylogenetic diversity, we calculated the standardized effect size of mean pairwise distances (ses.MPD) and the standard effect size of mean nearest taxon distances (ses.MNTD) in communities. For functional diversity, we calculated functional richness (FRic), functional evenness (FEve), functional divergence (FDiv), and the community-level weighted means (CWM) of trait values for diet, foraging stratum, and body mass, separately. From the 1950s to 2010s, species richness declined without significant trends. The ses.MNTD of Anatidae communities showed no clear trends. However, ses.MPD of Anatidae communities declined dramatically during this period. For functional diversity, functional evenness of diet, foraging stratum, body mass, and functional dispersion of diet, foraging stratum did not increase or decline significantly. However, functional evenness of all traits, functional richness, and functional dispersion of body mass showed declined trends. The basic phylogenetic diversity and species body mass of Anatidae communities declined significantly because of a declining trend in the relative independent branch of geese. This makes it more challenging for implement community recovery in the future. More attention in conservation biology should consider taxonomic diversity and asynchrony in phylogenetic and functional diversity.

Eiders, nutrients and eagles: Bottom-up and top-down population dynamics in a marine bird.

The main objective of this long-term study (1978-2016) was to find the underlying factors behind the declining trends of eider Somateria mollissima in the Baltic/Wadden Sea. Specifically, we aimed at quantifying the bottom-up effect of nutrients, through mussel stocks, on reproduction and abundance of eider, and the top-down effects caused by white-tailed eagle Haliaeetus albicilla predation. Bottom-up effects increase marine primary productivity with subsequent effects on food availability for a major mussel predator. Top-down effects may also regulate eider populations because during incubation female eiders are vulnerable to predation by eagles. Our structural equation modelling explained a large percentage of the variance in eider abundance. We conclude that the Baltic/Wadden Sea eider population was regulated directly by white-tailed sea eagle predation on incubating females and indirectly by the amount of nutrients in seawater affecting both mussel stocks and the breeding success of eiders, reflecting density dependence. These findings may explain the decreasing trend in the Baltic/Wadden Sea eider population during the last decades as an additive effect of top-down and bottom-up factors, and likely as an interaction between them.

Cuckoos use host egg number to choose host nests for parasitism.

To maximize their offspring success common cuckoo (Cuculus canorus) females should lay their eggs into host nests before incubation has begun. This ensures that the parasite chick hatches before all host chicks and can evict its foster siblings to monopolize host parental care. Many studies have demonstrated that most cuckoo eggs are indeed laid before the onset of host incubation. But cues used by female cuckoos to choose the right nest at the right time remain unclear. Here, we combine field observations with a field experiment to test whether female cuckoos use the number of eggs in the nests of their Oriental reed warbler (Acrocephalus orientalis) hosts to direct their choice. Over 8 years of field observations and 5 years of experiments, cuckoo females placed the majority of their eggs in nests with fewer than three host eggs, i.e. early in the laying sequence. For natural nests, the cuckoos may use information gleaned from the activity and behaviour of the host parents to make their choice. In our sets of experimental nests containing different numbers of model eggs, the vast majority of parasitism events occurred in nests containing a single egg. To our knowledge, this is the first field experiment showing that cuckoos choose host nests for parasitism based on the number of host eggs they contain. It appears that cuckoo females use the egg number to estimate the appropriate host nest stage for timely parasitism.

Model eggs fail to detect egg recognition in host populations after brood parasitism is relaxed.

BACKGROUND: Obligate brood parasites exert strong selective pressure on target hosts. In response, hosts typically evolve anti-parasitism strategies, of which egg recognition is one of the most efficient. Generally, host egg-recognition capacity is determined using model eggs. Previous studies have shown that some host species, which are capable of detecting parasite eggs, do not reject model eggs. However, it is unknown that whether the reaction to model eggs varies among distinct populations of the same host in relation to the degree of parasitism pressure. RESULTS: Here, we compared the rejection frequencies of model eggs and real eggs between mainland and island populations of the plain prinia (Prinia inornata), which are respectively sympatric and allopatric with their brood parasite, the common cuckoo (Cuculus canorus). Our results indicated that the mainland and island populations rejected real eggs at similar rates, but rejected model eggs, which were similar in size to real eggs but heavier, at significantly different rates: the island population rejected fewer model eggs, possibly because the rejection motivation of this population was lower due to absence of parasitism. CONCLUSIONS: Our results indicated that some factors affecting the decision to reject, such as rejection motivation, varied according to the degree of parasitism pressure, and thus influenced the frequency of egg rejection. Furthermore, our results suggested that model eggs should be used with caution in comparative studies of egg recognition abilities among species or populations subjected to different intensities of brood parasitism. That is, model eggs may fail to accurately detect egg recognition in host populations with little to no risk of parasitism.

Wing morphology, flight type and migration distance predict accumulated fuel load in birds.

Birds often accumulate large fat and protein reserves to fuel long-distance flights. While it is well known that species that fly the longest accumulate the largest amounts of fuel, considerable cross-species variation in fuel load is seen after controlling for overall migration distance. It remains unclear whether this variation can be explained by aerodynamic attributes of different species, despite obvious ecological and conservation implications. Here, we collected data on wing morphology, flight type, migration distance and fuel load from 213 European bird species and explored three questions: (1) does maximum fuel load relate to migration distance across species?; (2) does wing morphology, as described by wing aspect ratio and wing loading, influence maximum fuel load?; and (3) does flight type influence maximum fuel load? Our results indicate that maximum fuel load increases with migration across species, but residual variance is high. The latter variance is explained by aspect ratio and flight type, while wing loading and body mass explain little variance. Birds with slender wings accumulate less fuel than species with low wing aspect ratio when covering a similar migration distance. Continuously flapping species accumulate the largest amounts of fuel, followed by flapping and soaring species and flapping and gliding species, while the smallest fuel loads were observed in birds with passerine-type flight. These results highlight complex eco-evolutionary adaptations to migratory behaviour, pointing toward the importance of energy minimisation.

Heterospecific alarm-call recognition in two warbler hosts of common cuckoos.

Species facing similar selection pressures should recognize heterospecific alarm signals. However, no study has so far examined heterospecific alarm-call recognition in response to parasitism by cuckoos. In this study, we tested whether two sympatric host species of the common cuckoo Cuculus canorus, Oriental reed warbler Acrocephalus orientalis (ORW, main host), and black-browed reed warbler Acrocephalus bistrigiceps (BRW, rare host), could recognize each other's alarm calls in response to cuckoos. Dummies of common cuckoo (parasite) and Eurasian sparrowhawk Accipiter nisus (predator) were used to induce and record alarm calls of the two warbler species, respectively. In the conspecific alarm-call playback experiments, ORW responded more strongly to cuckoo alarm calls than to sparrowhawk alarm calls, while BRW responded less strongly to cuckoo alarm calls than to sparrowhawk alarm calls. In the heterospecific alarm-call playback experiments, both ORW and BRW responded less strongly to cuckoo alarm calls than sparrowhawk alarm calls. BRW seemed to learn the association between parasite-related alarm calls of the ORW and the cuckoo by observing the process of ORW attacking cuckoos. In contrast, alarm calls of BRW to cuckoos were rarely recorded in most cases. BRW with low parasite pressure still developed recognition of heterospecific parasite-related alarm call. Unintended receivers in the same community should recognize heterospecific alarm calls precisely to extract valuable information.

Predation and nutrients drive population declines in breeding waders.

Allee effects are defined as a decline in per capita fitness at low population density. We hypothesized that predation reduces population size of breeding waders and thereby the efficiency of predator deterrence, while total nitrogen through its effects on primary and secondary productivity increases population size. Therefore, nest predation could have negative consequences for population size because nest failure generally results in breeding dispersal and hence reduced local population density. To test these predictions, we recorded nest predation in five species of waders for 4,745 nests during 1987-2015 at the nature reserve Tipperne, Denmark. Predation rates were generally negatively related to conspecific and heterospecific population density, but positively related to overall population density of the entire wader community. Nest predation and population density were related to ground water level, management (grazing and mowing), and nutrients. High nest predation with a time lag of one year resulted in low overall breeding population density, while high nutrient levels resulted in higher population density. These two factors accounted for 86% of the variance in population size, presumably due to effects of nest predation on emigration, while nutrient levels increased the level of vegetation cover and the abundance of food in the surrounding brackish water. These findings are consistent with the hypothesis that predation may reduce population density through negative density dependence, while total nitrogen at adjacent shallow water may increase population size. Nest predation rates were reduced by high ground water level in March, grazing by cattle and mowing that affected access to and susceptibility of nests to predators. These effects can be managed to benefit breeding waders.

Cuckoos host range is associated positively with distribution range and negatively with evolutionary uniqueness.

The evolutionary distinctiveness (ED) score is a measure of phylogenetic isolation that quantifies the evolutionary uniqueness of a species. Here, we compared the ED score of parasitic and non-parasitic cuckoo species world-wide, to understand whether parental care or parasitism represents the largest amount of phylogenetic uniqueness. Next, we focused only on 46 cuckoo species characterized by brood parasitism with a known number of host species, and we explored the associations among ED score, number of host species and breeding range size for these species. We assessed these associations using phylogenetic generalized least squares (PGLS) models, taking into account the phylogenetic signal. Parasitic cuckoo species were not more unique in terms of ED than non-parasitic species. However, we found a significant negative association between the evolutionary uniqueness and host range and a positive correlation between the number of host species and range size of parasitic cuckoos, probably suggesting a passive sampling of hosts by parasitic species as the breeding range broadens. The findings of this study showed that more generalist brood parasites occupied very different positions in a phylogenetic tree, suggesting that they have evolved independently within the Cuculiformes order. Finally, we demonstrated that specialist cuckoo species also represent the most evolutionarily unique species in the order of Cuculiformes.

Effects of interspecific coexistence on laying date and clutch size in two closely related species of hole-nesting birds.

Coexistence between great tits Parus major and blue tits Cyanistes caeruleus, but also other hole-nesting taxa, constitutes a classic example of species co-occurrence resulting in potential interference and exploitation competition for food and for breeding and roosting sites. However, the spatial and temporal variations in coexistence and its consequences for competition remain poorly understood. We used an extensive database on reproduction in nest boxes by great and blue tits based on 87 study plots across Europe and Northern Africa during 1957-2012 for a total of 19,075 great tit and 16,729 blue tit clutches to assess correlative evidence for a relationship between laying date and clutch size, respectively, and density consistent with effects of intraspecific and interspecific competition. In an initial set of analyses, we statistically controlled for a suite of site-specific variables. We found evidence for an effect of intraspecific competition on blue tit laying date (later laying at higher density) and clutch size (smaller clutch size at higher density), but no evidence of significant effects of intraspecific competition in great tits, nor effects of interspecific competition for either species. To further control for site-specific variation caused by a range of potentially confounding variables, we compared means and variances in laying date and clutch size of great and blue tits among three categories of difference in density between the two species. We exploited the fact that means and variances are generally positively correlated. If interspecific competition occurs, we predicted a reduction in mean and an increase in variance in clutch size in great tit and blue tit when density of heterospecifics is higher than the density of conspecifics, and for intraspecific competition, this reduction would occur when density of conspecifics is higher than the density of heterospecifics. Such comparisons of temporal patterns of means and variances revealed evidence, for both species, consistent with intraspecific competition and to a smaller extent with interspecific competition. These findings suggest that competition associated with reproductive behaviour between blue and great tits is widespread, but also varies across large spatial and temporal scales.

Keeping eggs warm: thermal and developmental advantages for parasitic cuckoos of laying unusually thick-shelled eggs.

Obligate brood parasites have evolved unusually thick-shelled eggs, which are hypothesized to possess a variety of functions such as resistance to puncture ejection by their hosts. In this study, we tested the hypothesis that obligate brood parasites lay unusually thick-shelled eggs to retain more heat for the developing embryo and thus contribute to early hatching of parasite eggs. By doing so, we used an infrared thermal imaging system as a non-invasive method to quantify the temperature of eggshells of common cuckoos (Cuculus canorus) and their Oriental reed warbler (Acrocephalus orientalis) hosts in an experiment that artificially altered the duration of incubation. Our results showed that cuckoo eggshells had higher temperature than host eggs during incubation, but also less fluctuations in temperature during incubation disturbance. Therefore, there was a thermal and hence a developmental advantage for brood parasitic cuckoos of laying thick-shelled eggs, providing another possible explanation for the unusually thick-shelled eggs of obligate brood parasites and earlier hatching of cuckoo eggs compared to those of the host.

When a duck is not a duck; a new interdisciplinary synthesis for environmental radiation protection.

This consensus paper presents the results of a workshop held in Essen, Germany in September 2017, called to examine critically the current approach to radiological environmental protection. The meeting brought together participants from the field of low dose radiobiology and those working in radioecology. Both groups have a common aim of identifying radiation exposures and protecting populations and individuals from harmful effects of ionising radiation exposure, but rarely work closely together. A key question in radiobiology is to understand mechanisms triggered by low doses or dose rates, leading to adverse outcomes of individuals while in radioecology a key objective is to recognise when harm is occurring at the level of the ecosystem. The discussion provided a total of six strategic recommendations which would help to address these questions.

Ionizing radiation and taxonomic, functional and evolutionary diversity of bird communities.

Ionizing radiation from nuclear accidents at Chernobyl, Fukushima and elsewhere has reduced the abundance, species richness and diversity of ecosystems. Here we analyzed the taxonomic, functional and evolutionary diversity of bird communities in forested areas around Chernobyl. Species richness decreased with increasing radiation, mainly in 2007. Functional richness, but not functional evenness and divergence, decreased with increasing level of ionizing radiation. Evolutionary distinctiveness of bird communities was higher in areas with higher levels of ionizing radiation. Regression tree models revealed that species richness was higher in bird communities in areas with radiation levels lower than 0.7 µSv/h. In contrast, when radiation levels were higher than 16.67 µSv/h, bird species richness reached a minimum. Functional richness was affected by two variables: Forest cover and radiation level. Higher functional richness was found in bird communities in areas with forest cover lower than 50%. In the areas with forest cover higher than 50%, the functional richness was lower when radiation level was higher than 0.91 µSv/h. Finally, the average evolutionary distinctiveness of bird communities was higher in areas with forest cover exceeding 50%. These findings imply that level of ionizing radiation interacted with forest cover to affect species richness and its component parts, i.e. taxonomic, functional, and evolutionary diversity.

Partitioning within-species variance in behaviour to within- and between-population components for understanding evolution.

Phenotypes vary at multiple hierarchical levels, of which the interspecific variance is the primary focus of phylogenetic comparative studies. However, the evolutionary role of particular within-species variance components (between-population, between- or within-individual variances) remains neglected. Here, we partition the variance in an anti-predator behaviour, flight initiation distance (FID), and assess how its within- and between-population variance are related to life history, distribution, dispersal and habitat ecology. Although the composition of within-species variance in FID depended on the phylogeny, most variance occurred within populations. When accounting for allometry, density-dependence, uncertainty in the phylogenetic hypothesis and heterogeneity in data quality, within-population variance was significantly associated with habitat diversity and population size. Between-population variance was a significant predictor of natal dispersal, senescence and habitat diversity. Accordingly, not only species-specific mean values of a behavioural trait, but also its variance within and among populations can shape the evolutionary ecology of species.

Climate predicts which sex acts as helpers among cooperatively breeding bird species.

Among avian cooperative breeders, help in raising offspring is usually provided by males or by both sexes. Sex bias in helping should evolve in response to sex-specific ecological constraints on independent reproduction, with mate shortage for males and breeding vacancy shortage for each sex. Given that male-biased adult sex ratios are prevalent among birds, we predict that male-only helping mainly occurs in temperate species where fast population turnovers deriving from low adult annual survival allow all adult females to hold breeding vacancies, whereas some males overflow as helpers, and both-sex helping in tropical species where saturated habitats prevent not only males, but also females from breeding themselves. As expected, we found that across species, adult survival increased towards tropical zones and warmer climates, and higher adult survival tended to be associated with both-sex helping. Furthermore, sex bias in helping was predicted by latitude and ambient temperature. Our findings of demographic response of species to climate as a potential determinant of bias in helper sex uncover how ecological constraints operate to limit independent reproduction in sex-specific ways.