The effects of cities on quail (Coturnix coturnix) migration: a disturbing story of population connectivity, health, and ecography.

The increasing impact of human activities on ecosystems is provoking a profound and dangerous effect, particularly in wildlife. Examining the historical migration patterns of quail (Coturnix coturnix) offers a compelling case study to demonstrate the repercussions of human actions on biodiversity. Urbanization trends, where people gravitate toward mega-urban areas, amplify this effect. The proliferation of artificial urban ecosystems extends its influence across every biome, as human reliance on infrastructure and food sources alters ecological dynamics extensively. We examine European quail migrations pre- and post-World War II and in the present day. Our study concentrates on the Italian peninsula, investigating the historical and contemporary recovery of ringed quail populations. To comprehend changes in quail migration, we utilize trajectory analysis, open statistical data, and linear generalized models. We found that while human population and economic growth have shown a linear increase, quail recovery rates exhibit a U-shaped trajectory, and cereal and legume production displays an inverse U-shaped pattern. Generalized linear models have unveiled the significant influence of several key factors-time periods, cereal and legume production, and human demographics-on quail recovery rates. These factors closely correlate with the levels of urbanization observed across these timeframes. These insights underscore the profound impact of expanding human populations and the rise of mega-urbanization on ecosystem dynamics and services. As our planet becomes more urbanized, the pressure on ecosystems intensifies, highlighting the urgent need for concerted efforts directed toward conserving and revitalizing ecosystem integrity. Simultaneously, manage the needs and demands of burgeoning mega-urban areas. Achieving this balance is pivotal to ensuring sustainable coexistence between urban improvement and the preservation of our natural environment.

Eco-evolutionary drivers of avian migratory connectivity.

Migratory connectivity, reflecting the extent by which migrants tend to maintain their reciprocal positions in seasonal ranges, can assist in the conservation and management of mobile species, yet relevant drivers remain unclear. Taking advantage of an exceptionally large (~150,000 individuals, 83 species) and more-than-a-century-long dataset of bird ringing encounters, we investigated eco-evolutionary drivers of migratory connectivity in both short- and long-distance Afro-Palearctic migratory birds. Connectivity was strongly associated with geographical proxies of migration costs and was weakly influenced by biological traits and phylogeny, suggesting the evolutionary lability of migratory behaviour. The large intraspecific variability in avian migration strategies, through which most species geographically split into distinct migratory populations, explained why most of them were significantly connected. By unravelling key determinants of migratory connectivity, our study improves knowledge about the resilience of avian migrants to ecological perturbations, providing a critical tool to inform transboundary conservation and management strategies at the population level.

A deepening understanding of animal culture suggests lessons for conservation.

A key goal of conservation is to protect biodiversity by supporting the long-term persistence of viable, natural populations of wild species. Conservation practice has long been guided by genetic, ecological and demographic indicators of risk. Emerging evidence of animal culture across diverse taxa and its role as a driver of evolutionary diversification, population structure and demographic processes may be essential for augmenting these conventional conservation approaches and decision-making. Animal culture was the focus of a ground-breaking resolution under the Convention on the Conservation of Migratory Species of Wild Animals (CMS), an international treaty operating under the UN Environment Programme. Here, we synthesize existing evidence to demonstrate how social learning and animal culture interact with processes important to conservation management. Specifically, we explore how social learning might influence population viability and be an important resource in response to anthropogenic change, and provide examples of how it can result in phenotypically distinct units with different, socially learnt behavioural strategies. While identifying culture and social learning can be challenging, indirect identification and parsimonious inferences may be informative. Finally, we identify relevant methodologies and provide a framework for viewing behavioural data through a cultural lens which might provide new insights for conservation management.

Prevalence and genetic diversity of avian haemosporidian parasites in wild bird species of the order Columbiformes.

Diseases can play a role in species decline. Among them, haemosporidian parasites, vector-transmitted protozoan parasites, are known to constitute a risk for different avian species. However, the magnitude of haemosporidian infection in wild columbiform birds, including strongly decreasing European turtle doves, is largely unknown. We examined the prevalence and diversity of haemosporidian parasites Plasmodium, Leucocytozoon and subgenera Haemoproteus and Parahaemoproteus in six species of the order Columbiformes during breeding season and migration by applying nested PCR, one-step multiplex PCR assay and microscopy. We detected infections in 109 of the 259 screened individuals (42%), including 15 distinct haemosporidian mitochondrial cytochrome b lineages, representing five H. (Haemoproteus), two H. (Parahaemoproteus), five Leucocytozoon and three Plasmodium lineages. Five of these lineages have never been described before. We discriminated between single and mixed infections and determined host species-specific prevalence for each parasite genus. Observed differences among sampled host species are discussed with reference to behavioural characteristics, including nesting and migration strategy. Our results support previous suggestions that migratory birds have a higher prevalence and diversity of blood parasites than resident or short-distance migratory species. A phylogenetic reconstruction provided evidence for H. (Haemoproteus) as well as H. (Parahaemoproteus) infections in columbiform birds. Based on microscopic examination, we quantified parasitemia, indicating the probability of negative effects on the host. This study provides a large-scale baseline description of haemosporidian infections of wild birds belonging to the order Columbiformes sampled in the northern hemisphere. The results enable the monitoring of future changes in parasite transmission areas, distribution and diversity associated with global change, posing a potential risk for declining avian species as the European turtle dove.

Crimean-Congo Hemorrhagic Fever Virus Genome in Tick from Migratory Bird, Italy.

We detected Crimean-Congo hemorrhagic fever virus in a Hyalomma rufipes nymph collected from a whinchat (Saxicola rubetra) on the island of Ventotene in April 2017. Partial genome sequences suggest the virus originated in Africa. Detection of the genome of this virus in Italy confirms its potential dispersion through migratory birds.

Clock gene polymorphism, migratory behaviour and geographic distribution: a comparative study of trans-Saharan migratory birds.

Migratory behaviour is controlled by endogenous circannual rhythms that are synchronized by external cues, such as photoperiod. Investigations on the genetic basis of circannual rhythmicity in vertebrates have highlighted that variation at candidate 'circadian clock' genes may play a major role in regulating photoperiodic responses and timing of life cycle events, such as reproduction and migration. In this comparative study of 23 trans-Saharan migratory bird species, we investigated the relationships between species-level genetic variation at two candidate genes, Clock and Adcyap1, and species' traits related to migration and geographic distribution, including timing of spring migration across the Mediterranean Sea, migration distance and breeding latitude. Consistently with previous evidence showing latitudinal clines in 'circadian clock' genotype frequencies, Clock allele size increased with breeding latitude across species. However, early- and late-migrating species had similar Clock allele size. Species migrating over longer distances, showing delayed spring migration and smaller phenotypic variance in spring migration timing, had significantly reduced Clock (but not Adcyap1) gene diversity. Phylogenetic confirmatory path analysis suggested that migration date and distance were the most important variables directly affecting Clock gene diversity. Hence, our study supports the hypothesis that Clock allele size increases poleward as a consequence of adaptation to the photoperiodic regime of the breeding areas. Moreover, we show that long-distance migration is associated with lower Clock diversity, coherently with strong stabilizing selection acting on timing of life cycle events in long-distance migratory species, likely resulting from the time constraints imposed by late spring migration.

Migratory connectivity and effects of winter temperatures on migratory behaviour of the European robin Erithacus rubecula: a continent-wide analysis.

Many partially migratory species show phenotypically divergent populations in terms of migratory behaviour, with climate hypothesized to be a major driver of such variability through its differential effects on sedentary and migratory individuals. Based on long-term (1947-2011) bird ringing data, we analysed phenotypic differentiation of migratory behaviour among populations of the European robin Erithacus rubecula across Europe. We showed that clusters of populations sharing breeding and wintering ranges varied from partial (British Isles and Western Europe, NW cluster) to completely migratory (Scandinavia and north-eastern Europe, NE cluster). Distance migrated by birds of the NE (but not of the NW) cluster decreased through time because of a north-eastwards shift in the wintering grounds. Moreover, when winter temperatures in the breeding areas were cold, individuals from the NE cluster also migrated longer distances, while those of the NW cluster moved over shorter distances. Climatic conditions may therefore affect migratory behaviour of robins, although large geographical variation in response to climate seems to exist.

Polymorphism at the Clock gene predicts phenology of long-distance migration in birds.

Dissecting phenotypic variance in life history traits into its genetic and environmental components is at the focus of evolutionary studies and of pivotal importance to identify the mechanisms and predict the consequences of human-driven environmental change. The timing of recurrent life history events (phenology) is under strong selection, but the study of the genes that control potential environmental canalization in phenological traits is at its infancy. Candidate genes for circadian behaviour entrained by photoperiod have been screened as potential controllers of phenological variation of breeding and moult in birds, with inconsistent results. Despite photoperiodic control of migration is well established, no study has reported on migration phenology in relation to polymorphism at candidate genes in birds. We analysed variation in spring migration dates within four trans-Saharan migratory species (Luscinia megarhynchos; Ficedula hypoleuca; Anthus trivialis; Saxicola rubetra) at a Mediterranean island in relation to Clock and Adcyap1 polymorphism. Individuals with larger number of glutamine residues in the poly-Q region of Clock gene migrated significantly later in one or, respectively, two species depending on sex and whether the within-individual mean length or the length of the longer Clock allele was considered. The results hinted at dominance of the longer Clock allele. No significant evidence for migration date to covary with Adcyap1 polymorphism emerged. This is the first evidence that migration phenology is associated with Clock in birds. This finding is important for evolutionary studies of migration and sheds light on the mechanisms that drive bird phenological changes and population trends in response to climate change.

Avian Haemosporidians Infecting Short- and Long-Distance Migratory Old World Flycatcher Species and the Variation in Parasitaemia After Endurance Flights.

PURPOSE: Avian haemosporidians are widespread parasites, occurring in many bird families and causing pathologies ranging from rather benign infections to highly virulent diseases. The state of knowledge about lineage-specific intensities of haemosporidian infection (i.e., parasitaemia) is mainly based on infection experiments conducted under laboratory conditions. The levels and range of parasitaemia in natural host-parasite associations as well as their influencing factor remain largely unexplored. METHODS: Thus, we explored the parasitaemia of four songbird species (i.e., European Robins, Black and Common Redstarts and Whinchats) during migration by screening individuals upon landing on an insular passage site after extensive endurance flights to (1) describe their natural host-parasite associations, (2) quantify parasitaemia and (3) explore potential host- and parasite-related factors influencing parasitaemia. RESULTS: We found 68% of Whinchats to be infected with haemosporidians, which is more frequent than any other of the studied host species (30-34%). Furthermore, we confirmed that parasitaemia of Haemoproteus infections was higher than average Plasmodium infections. Median parasitaemia levels were rather low (parasite cells in 0.01% of hosts' red blood cells) and varied largely among the different parasite lineages. However, we found four individuals hosting infections with parasitaemia higher than typical chronic infections. CONCLUSIONS: Based on the known transmission areas of the respective lineages, we argue that these higher intensity infections might be relapses of consisting infections rather than acute phases of recent primary infections.

Reconstructing migratory network nodes to improve environmental management and conservation decisions: A case study of the common quail Coturnix coturnix as a biosensor.

Public health risks are related to environmental management practices, and are essential to the understanding of ecosystem dynamics. The expansion of urbanized areas affects migratory bird networks and can impact the population sizes of migratory species and, in parallel, increase the risk of diseases carried by migratory species spreading into urban areas. Taking the European population of the common quail as a case study, we reconstructed its migratory network between Europe and the Maghreb using quail recoveries from the Italian Bird Ringing Scheme. We showed that soils at the central European migratory node have been degraded through urbanization and reforestation, decreasing the number of successful migrations over this area. Conceptual models incorporating the One Health approach, natural and social capital, landscape infrastructure, and ecosystem services could explain the relationships between climate warming and extensively developed, intensively developed, and urbanized ecosystems. Failed migratory flights of quail over central Europe highlight not only the problems of infrastructure design and their impact on ecosystems services, but also on One Health parameters. The damage to migratory network nodes presents a global threat to biodiversity and can increase the spread of diseases. To address this challenge we propose: i) improvements in land quality; ii) transboundary migration monitoring programmes; and iii) management plans for migratory birds - the overall aim being to optimize infrastructures to improve the quality of human life. Lessons drawn from a better understanding of the migratory networks of quail in relation to different ecosystems provide useful tools to improve infrastructure management and political decision making processes.

Modelling the timing of migration of a partial migrant bird using ringing and observation data: a case study with the Song Thrush in Italy.

BACKGROUND: The study of the timing of migration is fundamental to the understanding of the ecology of many bird species and their response to climate change, and it has important conservation and management implications e.g., for assessing the hunting seasons according to the EU Directive 2009/147/EC (Birds Directive). METHODS: We developed a new method for the analysis of ringing data (both first capture and re-encounters) and citizen science observations, to assess the timing of pre- and post-nuptial migration of birds. This method was tested on the Song Thrush Turdus philomelos, using i) the Bird Ringing Database hosted by the ISPRA Italian Ringing Centre from the whole Italian peninsula, the three closest large islands (Sicily, Sardinia and Corsica), and Canton Ticino (Switzerland) and ii) the eBird data for the same study area. RESULTS: The results from both datasets consistently showed that pre-nuptial migration starts during the first 10-day period of January (Jan 1) in some central and southern areas of the Italian peninsula, in central Sicily, southern Sardinia, and Corsica. The onset of migration occurs on Jan 2 in the rest of central and southern Italy, Sicily and Sardinia, and western Liguria, while it starts later in the north-eastern Alps, up to Mar 3. The end of post-nuptial migration is more synchronous, occurring on Nov 1 across most of Italy, slightly earlier (Oct 3) in northern Italy and later (Nov 2) in Sicily. The uncertainty of the estimated dates was < 2 days in most cases. CONCLUSION: This method represents a novel and valuable tool for the analyses of the timing of migration using ringing and citizen science data and provides an important contribution to the Key Concepts Document of the EU Birds Directive, where migration timings are considered and used to define the hunting period of birds.

Short- and long-distance avian migrants differ in exercise endurance but not aerobic capacity.

BACKGROUND: Migratory birds differ markedly in their migration strategies, particularly those performing short- versus long-distance migrations. In preparation for migration, all birds undergo physiological and morphological modifications including enlargement of fat stores and pectoral muscles to fuel and power their flights, as well as cardiovascular and biochemical adjustments that improve lipid and oxygen delivery and uptake by flight muscles. While the magnitude of these changes varies in relation to migration strategy, the consequence of these variations on aerobic performance is unknown. We tested whether the aerobic performance of four Old-world flycatcher species (Muscicapidae) varied according to migration strategy by comparing minimum resting metabolic rates (RMRmin), exercise-induced maximum metabolic rates (MMR), and exercise endurance times of short-distance and long-distance migratory birds. RESULTS: As expected, RMRmin did not vary between short-distance and long-distance migrants but differed between the species within a migration strategy and between sexes. Unexpectedly, MMR did not vary with migration strategy, but MMR and blood haemoglobin content were positively related among the birds tested. Exercise endurance times differed substantially between migration strategies with long-distance migrants sustaining exercise for > 60% longer than short-distance migrants. Blood haemoglobin content had a significant positive effect on endurance among all birds examined. CONCLUSIONS: The lack of difference in RMRmin and MMR between long- and short-distance migrants during this stage of migration suggests that the attributes favouring the greater aerobic endurance of long-distance migrants did not come at the expense of increased maintenance costs or require greater aerobic capacity.

West Nile and Usutu Virus Introduction via Migratory Birds: A Retrospective Analysis in Italy.

The actual contribution of migratory birds in spreading West Nile (WNV) and Usutu virus (USUV) across Europe and from Africa to old countries is still controversial. In this study, we reported the results of molecular and serological surveys on migrating birds sampled during peaks of spring and autumn migration at 11 Italian sites located along important flyways, from 2012 to 2014. A total of 1335 specimens made of individual or pooled sera, and organs from 275 dead birds were tested for WNV and USUV RNA by real time PCR (RT-PCR). Furthermore, sera were tested by serum neutralization assay for detecting WNV and USUV neutralizing antibodies. Molecular tests detected WNV lineage 2 RNA in a pool made of three Song Thrush (Turdus philomelos) sera sampled in autumn, and lineage 1 in kidneys of six trans-Saharan birds sampled in spring. Neutralizing antibodies against WNV and USUV were found in 5.80% (n = 72; 17 bird species) and 0.32% (n = 4; 4 bird species) of the tested sera, respectively. Our results do not exclude the role of migratory birds as potential spreaders of WNV and USUV from Africa and Central Europe to Mediterranean areas and highlight the importance of a more extensive active surveillance of zoonotic viruses.

Direct and Indirect Role of Migratory Birds in Spreading CCHFV and WNV: A Multidisciplinary Study on Three Stop-Over Islands in Italy.

The annual movements of migratory birds can contribute to the spread of African ticks and tick-borne pathogens of potential public health concern across Europe. The aim of the study was to investigate their role in the possible introduction of African ticks and tick-borne pathogens into European countries during spring migration. A total of 2344 ticks were collected during three spring seasons from 1079 birds captured on three Italian stop-over islands during their northbound migration. Once identified, each tick was tested by RT-PCR for the presence of Crimean-Congo hemorrhagic fever (CCHFV), West Nile (WNV), and Usutu (USUV) viruses. Moreover, carcasses of birds found dead were collected and tested for the possible presence of WNV and USUV. Results confirmed a higher contribution of trans-Saharan migrants compared to intra-Palearctic ones and the prevalence of African tick species in the sample. CCHFV was detected for the second time in Italy in a Hyalomma rufipes, and WNV was found in two ticks of the same genus, all carried by trans-Saharan birds. WNV lineage 1 was also found in the organs of a Garden warbler. These results confirm the role of migratory birds in carrying African ticks, as well as viruses of zoonotic importance, from Africa into Europe.

The genome sequence of the European nightjar, Caprimulgus europaeus (Linnaeus, 1758).

We present a genome assembly from an individual female Caprimulgus europaeus (the European nightjar; Chordata; Aves; Caprimulgiformes; Caprimulgidae). The genome sequence is 1,178 megabases in span. The majority of the assembly (99.33%) is scaffolded into 37 chromosomal pseudomolecules, including the W and Z sex chromosomes.

Comparing population trend estimates of migratory birds from breeding censuses and capture data at a spring migration bottleneck.

Europe has a well-established network of breeding bird monitoring that is used to produce supranational indices of population trends for many species. However, a comparison of breeding bird censuses with other methods may be beneficial to confirm the validity of such indices. The aim of this study was to assess the value of standardized capture data of migratory birds at migration bottlenecks as an indicator of the effective breeding populations. One limitation to this method is that several populations are co-occurring at these bottlenecks and their catchment areas need to be clearly identified to allow extrapolation of population indices. Here, we used standardized trends in capture numbers of 30 species on the island of Ponza, a migration bottleneck in the central Mediterranean, and compared them to population trends estimated in the putative catchment breeding areas between 2005 and 2016. The catchment areas were identified through the analysis of ring recoveries during the breeding season of birds passing through Ponza. Our results show an agreement between the population trends observed on Ponza and those in the breeding areas in 15 out of 30 species. The correlations were strongest in species with a more robust definition of the catchment areas, that is, species with more than 10 recoveries, and for which the recoveries were most likely of breeding birds. The main reason for disagreement between the two indices in the remaining species might be related to different intensity of sampling in different areas. This issue can be solved by further developing monitoring projects in underrepresented countries, as well as by intensifying monitoring through ringing, both in the breeding grounds and at migration bottlenecks. These results show that spring migration monitoring at bottlenecks has the potential to provide a valuable complement and an independent control of breeding bird surveys, allowing raising early warnings of population declines and contributing to their conservation.

Body fat influences departure from stopover sites in migratory birds: evidence from whole-island telemetry.

Migration remains one of the great mysteries of animal life. Small migratory birds rely on refuelling stopovers after crossing ecological barriers such as deserts or seas. Previous studies have suggested that fuel reserves may determine stopover duration but this hypothesis could not be tested because of methodological limitations. Here, we provide evidence that subcutaneous fat stores determine stopover duration by measuring the permanence of migratory garden warblers (Sylvia borin) on a small Mediterranean island during spring migration with telemetry methods. Garden warblers with large amounts of fat stores departed the island significantly sooner than lean birds. All except one fat bird left the island on the same evening after capture, with a mean total stopover estimate of 8.8 hours. In contrast, the mean estimated total stopover duration of lean birds was 41.3 hours. To our knowledge, this is the first study that measures the true minimum stopover duration of a songbird during migration.

Recent phenological shifts of migratory birds at a Mediterranean spring stopover site: Species wintering in the Sahel advance passage more than tropical winterers.

Spring migration phenology is shifting towards earlier dates as a response to climate change in many bird species. However, the patterns of change might not be the same for all species, populations, sex and age classes. In particular, patterns of change could differ between species with different ecology. We analyzed 18 years of standardized bird capture data at a spring stopover site on the island of Ponza, Italy, to determine species-specific rates of phenological change for 30 species following the crossing of the Mediterranean Sea. The advancement of spring passage was more pronounced in species wintering in Northern Africa (i.e. short-distance migrants) and in the Sahel zone. Only males from species wintering further South in the forests of central Africa advanced their passage, with no effect on the overall peak date of passage of the species. The migration window on Ponza broadened in many species, suggesting that early migrants within a species are advancing their migration more than late migrants. These data suggest that the cues available to the birds to adjust departure might be changing at different rates depending on wintering location and habitat, or that early migrants of different species might be responding differently to changing conditions along the route. However, more data on departure time from the wintering areas are required to understand the mechanisms underlying such phenological changes.

Evolution of chain migration in an aerial insectivorous bird, the common swift Apus apus.

Spectacular long-distance migration has evolved repeatedly in animals enabling exploration of resources separated in time and space. In birds, these patterns are largely driven by seasonality, cost of migration, and asymmetries in competition leading most often to leapfrog migration, where northern breeding populations winter furthest to the south. Here, we show that the highly aerial common swift Apus apus, spending the nonbreeding period on the wing, instead exhibits a rarely found chain migration pattern, where the most southern breeding populations in Europe migrate to wintering areas furthest to the south in Africa, whereas the northern populations winter to the north. The swifts concentrated in three major areas in sub-Saharan Africa during the nonbreeding period, with substantial overlap of nearby breeding populations. We found that the southern breeding swifts were larger, raised more young, and arrived to the wintering areas with higher seasonal variation in greenness (Normalized Difference Vegetation Index) earlier than the northern breeding swifts. This unusual chain migration pattern in common swifts is largely driven by differential annual timing and we suggest it evolves by prior occupancy and dominance by size in the breeding quarters and by prior occupancy combined with diffuse competition in the winter.

Assessing the role of migratory birds in the introduction of ticks and tick-borne pathogens from African countries: An Italian experience.

The continuous flow of billions of birds between Africa and Europe creates an "ecological bridge" between physically remote areas. Migratory birds fly south from their breeding grounds during late summer/fall and fly back in spring. These movements regulate the spread of internal and external parasites, as well as pathogens of potential public health concern. The aim of the present study was to investigate the possible introduction of exotic tick species and tick-borne pathogens into Europe via migratory birds. At the bird observatory of Ventotene island (Italy), 443 feeding ticks were collected from 249 birds captured and ringed during their northbound migration in spring 2013. Each tick was identified by morphological and molecular methods and then tested for bacterial and viral pathogens: Borrelia burgdorferi s.l., Rickettsia spp., Ehrlichia ruminantium and Coxiella burnetii, Crimean Congo haemorrhagic fever virus (CCHFV) and Flavivirus. Morphological and molecular identification confirmed Hyalomma rufipes as the most abundant species among the collected arthropods (366/443; 82.6%) followed by Hyalomma marginatum (10/433; 2.3%). Rickettsia aeschlimannii was identified in 158 ticks, while one engorged Amblyomma variegatum nymph was infected with Rickettsia africae. The other bacteria were not detected in any specimen. Among viruses, RNA belonging to West Nile virus and other Flavivirus were detected whereas all ticks were negative for CCHFV RNA. These results confirm how migratory birds play a role in carrying Rickettsia-infected ticks, as well as viruses of zoonotic importance, from Africa into Europe. To what extent tick species are capable of establishing a permanent population once introduced in naïve areas, is far from defined and deserve further investigation.