European Birds Research Articles

Detection ranges of forest bird vocalisations: guidelines for passive acoustic monitoring

Passive acoustic monitoring has proven to have many advantages for monitoring efforts and research activities. However, there are considerations to be taken into account regarding the placement of autonomous sound recorders. Detection ranges differ among species and in response to variable conditions such as weather or the location of vocalising animals. It is thus important to the success of a research project to understand, with a certain degree of confidence, the distances at which birds might be detected. In two types of forests in Poland, we played back the vocalisations of 31 species of European forest birds exemplifying different singing characteristics. Based on recordings obtained along a 500-m transect, we estimated the probability of detection and maximum detection distance of each vocalisation. We broadcasted the recording at three heights of singing and repeated playbacks three times during the breeding season to evaluate the effect of vegetation growth. Our results revealed that environmental and meteorological factors had a significant influence on both detection probability and maximum detection distances. This work provides comprehensive measurements of detection distance for 31 bird species and can be used to plan passive acoustic monitoring research in Europe, taking into account species traits and individual characteristics of the study area.

Design and preparation of a thematic showcase on alien species at the Museum of Zoology of the University of Catania (Italy)

As part of the Interreg FAST (Fight Alien Species Transborder) project, in which the Zoology Museum of the University of Catania (Italy) is involved, in the Museum main room a thematic exhibition relating to alien species, also equipped with interactive supports, has been planned. This action is part of the communication strategy of the aforementioned project, providing for the promotion of Citizen Science initiatives that bring all citizens closer to this problem, increasing their awareness and participation. Some of the most representative and iconic animals linked to this issue will be exhibited within this space. Visitors will be able to observe both vertebrates such as the Nutria (Myocastor coypus Molina, 1782) or the American pond turtle (Trachemys scripta Thunberg in Schoepff, 1792), and invertebrates such as the Red palm weevil (Rhynchophorus ferrugineus Olivier, 1790) or the Louisiana red crayfish (Procambarus clarkii Girard, 1852). Also, a large poster illustrating all the species of alien birds in Europe will complete the exhibition, which will also be accompanied by information panels in Italian and English regarding not only the animals on display but also the aims and objectives of the FAST project. Adjacent to the exhibition space an 80-inch touch screen monitor will be installed with a software, also for children, for the recognition of native and alien species, accompanied by information collected in a database on alien invasive species present in Sicily and in Malta.

Effect of 2020-21 and 2021-22 Highly Pathogenic Avian Influenza H5 Epidemics on Wild Birds, the Netherlands.

The number of highly pathogenic avian influenza (HPAI) H5-related infections and deaths of wild birds in Europe was high during October 1, 2020-September 30, 2022. To quantify deaths among wild species groups with known susceptibility for HPAI H5 during those epidemics, we collected and recorded mortality data of wild birds in the Netherlands. HPAI virus infection was reported in 51 bird species. The species with the highest numbers of reported dead and infected birds varied per epidemic year; in 2020-21, they were within the Anatidae family, in particular barnacle geese (Branta leucopsis) and in 2021-22, they were within the sea bird group, particularly Sandwich terns (Thalasseus sandvicensis) and northern gannet (Morus bassanus). Because of the difficulty of anticipating and modeling the future trends of HPAI among wild birds, we recommend monitoring live and dead wild birds as a tool for surveillance of the changing dynamics of HPAI.

Improving national bird population estimates in Europe: insights from comparisons with atlas abundance data

Summary In Europe, population estimates of breeding birds are produced nationally and are periodically compiled at EU or pan-European scales. Until now, no other source was available to explore the robustness of these estimates. In this study, we compared population sizes reported in the latest edition of the European Red List of Birds (ERLoB) with those produced using data from the second European Breeding Bird Atlas (EBBA2) to assess their consistency and determine parameters behind variability in population estimates that deserve further attention in the future. In general, European population estimates derived from summing local abundance data from EBBA2 were similar to those obtained from ERLoB, although for some species they differed considerably, particularly in those distributed mainly in southern Europe. National population estimates from EBBA2 also did not differ markedly from those in ERLoB. However, we found that EBBA2 provided larger national population sizes than ERLoB for widespread species, suggesting that spatial information is more relevant for properly assessing their population size than for localised species. Our analysis also showed that, in general, population estimates based on robust methodological protocols (e.g. complete counts, statistical inference) contributed to reducing differences between ERLoB and EBBA2 values. Interestingly, EBBA2 and ERLoB estimates were quite similar for species classified in Europe as “Threatened” or “Near Threatened”, whereas the values for “Least Concern” species were consistently different between these two sources. Our results indicate which type of species would benefit from additional efforts to improve national population estimates and their consistency across countries, issues that are of paramount importance for guiding conservation strategies in Europe.

Agroforestry supports high bird diversity in European farmland

Intensification and homogenization of agricultural landscapes have led to a strong decline in European farmland birds. Agroforestry systems, which were widespread in the past, are regaining attention as they could return structural heterogeneity to agricultural landscapes. However, few studies focus on the effects of such systems on biodiversity and especially bird diversity. We hypothesized that agroforestry systems host a higher alpha and beta diversity of birds compared to open agriculture as well as distinct bird communities. Moreover, we expected that bird communities in temperate Europe and the Mediterranean are differently affected by agroforestry systems. In this study, we assessed breeding bird diversity via audio recordings in nineteen mature agroforestry plots, comprising both silvoarable and silvopastoral systems distributed across seven countries in temperate and Mediterranean Europe. For comparison, bird diversity was also assessed in nearby open agricultural land, forests, and orchards. Bird species richness in agroforestry was more than doubled compared to open agricultural land and similar to the diversity found in forests and orchards. Community composition and within-habitat beta diversity differed between the habitat types and between European regions. While temperate agroforestry systems hosted generalist and woody habitat species, bird communities in Mediterranean agroforestry were composed of species from both open and woody habitats. Beta diversity was significantly higher in agroforestry than in open agriculture in temperate systems but not in the Mediterranean. Our study demonstrates that agroforestry systems represent a valuable habitat for breeding birds in European agricultural landscapes. A wider adoption of these systems could thus contribute to halting and reversing the decline in bird diversity, especially in temperate agricultural landscapes.

Genetic distinctiveness of an endangered falcon: Implications for conservation in Europe.

In the Falconidae, the genus Falco comprises species of large birds of prey with wide distribution worldwide. However, the European lanner falcon Falco biarmicus feldeggii is rapidly heading for global extinction following a dramatic decline caused by anthropogenic interference. Conservation projects are currently underway with the main purpose of increasing its population size in the Mediterranean basin through captive breeding and release of birds into the wild. To support the projects, and strengthen the legitimacy of conservation efforts consistently with the Evolutionary Significant Unit concept, we explored the possibility of characterising the gene pool of the European lanner and reliably distinguishing it from other falcon taxa inhabiting the Mediterranean area, which show morphological and genetic similarities. To address the issue, we examined genetic variability at the nuclear level through the analysis of 12 neutral Short Tandem Repeat loci, and, for the first time in these taxa, two single-copy functional genes, coding for the brain-derived neurotrophic factor precursor and the oocyte maturation factor, respectively. The second exon of the major histocompatibility complex class II B gene was also investigated. Additionally, to frame our data with previously published data, we assess variation at the mitochondrial level by sequencing portions of the cytochrome b, 12S rRNA gene, and the control region. Our results showed that the European lanner is highly distinct from other falcon taxa, as revealed by nuclear, but not by mitochondrial DNA. We discuss our findings focusing on their implications for the preservation of this highly endangered European bird, and highlighted the critical role of genetic information in planning and monitoring concrete interventions.

Seed Dispersal by an Invasive Exotic Bird in Europe

Seed Dispersal by an Invasive Exotic Bird in Europe

New molecular data help clarify the taxonomy of Central European avian Dicrocoeliidae Looss, 1899 (Trematoda: Plagiorchiida)

Avian Dicrocoeliidae are difficult to identify, even in their adult stages. Molecular analyses have been conducted for only a few species, complicating the accurate identification of juvenile stages. The taxonomy of the family is unresolved, and the status of many dicrocoeliid species is uncertain. Sequences of nuclear and mitochondrial DNA loci of Central European avian Dicrocoeliidae were generated and analyzed. These included representatives of the genera Lyperosomum, Platynosomum, Stromitrema, Brachylecithum, Brachydistomum, and Lutztrema. All the sequences were obtained from morphologically identified adult specimens of dicrocoeliids isolated from avian hosts. Molecular support was obtained to validate Lyperosomum turdia, confirm the rejection of Lyperosomum dujardini and Lyperosomum alagesi, and resurrect Lyperosomum longicauda and Lyperosomum collurionis. The identity of European Platynosomum illiciens from avian hosts with American vouchers of the same species from avian and mammalian hosts was confirmed. Brachylecithum fringillae is not considered valid; the individuals that matched its diagnosis were subadult Brachydistomum ventricosum. Descriptions and comparative data for five new species are provided. These are Lyperosomum hirundinis sp. n., Lyperosomum tenori sp. n., Lyperosomum atricapillae sp. n., Stromitrema acrocephali sp. n., and Lutztrema atricapillae sp. n.. Based on the molecular data, suggestions are provided regarding the validity of dicrocoeliid species that parasitize Central European birds. Further research should address the polyphyletic status of Brachylecithum.

Is Eriocnemis luciani meridae (Aves: Trochilidae) a diagnosable taxon and does it come from Venezuela, with remarks on the collectors Salomn Briceo and Walther Frederick Henninger.

Eriocnemis luciani meridae was originally described from a single specimen collected in the late 19th century in western Venezuela. Subsequently a second specimen of E. luciani, also labelled Venezuela, has been taken as additional proof for a highly disjunct population of this hummingbird, which otherwise ranges from southwest Colombia to southern Peru (taxonomy-dependent). Eriocnemis l. meridae has been accepted by all of the global checklists of birds, but has been routinely ignored by Venezuelan sources. In an effort to resolve this dichotomy of treatment, we re-examined the specimens plumage in comparison with relevant material in two major European bird collections. We found that the characters used to erect E. l. meridae are only doubtfully or weakly expressed in the holotype and appear invisible in the Ohio specimen, but both are clearly referrable to the species E. luciani. Evidence that the second specimen was definitely collected in Venezuela is weak and its overall provenance is unclear. In contrast, an extensive historical investigation of the relevant collectors indicates that the holotype does appear to have been taken in Venezuela, although perhaps not in the precise locality indicated for it. This leaves an unusual situation whereby we consider the case for a separate Venezuelan endemic taxon to be unproven, but there is no incontrovertible reason to exclude the species from the countrys avifauna; according to recent niche modelling data it is best searched for in the Sierra Nevada of Mrida state. In contrast, a second subspecies of E. luciani, E. l. baptistae, described by the same authors as endemic to part of western Ecuador is, according to our reappraisal, clearly diagnosable and is upheld.

A systematic review on the effectiveness of crop architecture-related in-field measures for promoting ground-breeding farmland birds

Ground-breeding farmland birds are disproportionately negatively affected by the impact of agricultural intensification and mostly still dramatically declining in Europe. One main reason for this decline is the lack of suitable nesting sites. While some off-field measures showed positive effects on bird productivity, we are currently not achieving sufficient conservation with these, so that supplementing them with in-field measures seems inevitable. Over the past years, several in-field measures have been developed aiming at providing suitable nesting sites in crop fields by manipulating the crop architecture, i.e., the density and/or height of the crops. However, we currently lack an overview on what has been tested and resulted in stabilizing population of ground-breeding farmland birds. In this systematic review, we provide a qualitative assessment of current knowledge and knowledge gaps on such in-field measures and their effects on European crop-breeding farmland birds. In doing so, we accounted for specific birds’ requirements on their breeding habitat. We found only very few studies on the effectiveness of crop architecture-related in-field measures on ground-breeding farmland birds. Knowledge gaps exist for effects on individual species in general, their reproduction (rather than population density), the influence of landscape and local contexts on the effectiveness, and the optimal spatial arrangement of measures to maximize their efficiency. This shows an urgent need for more research on a holistic scale. However, the few existing studies suggest, that there is a high potential for crop architecture-related in-field measures to promote ground-breeding farmland birds, and thus bring them back as ‘agricultural by-products’.

Urban landscape organization is associated with species-specific traits in European birds

Urbanization is one of the main current drivers of the global biodiversity loss. Cities are usually developed in a gradient between land-sharing (low density housing with small and fragmented green areas) and land-sparing areas (high density housing with large and non-fragmented green patches) depending on the spatial organization of urban attributes. Previous studies have indicated differences in biodiversity between these two urban development types, but mechanisms underlying these differences are inadequately understood. In this context, the landscape features of each urban development type may select for organisms with specific traits. To analyze it, we quantified birds in 9 European cities during the breeding and wintering season, collected species-specific traits and performed Bayesian comparative analyses. We found that birds living in land-sparing areas had a higher reproductive investment and a higher nesting specialization than birds living in land-sharing areas during the breeding season. Typical birds from land-sparing urban areas during winter are fast-lived species. Our results indicate that urban development type could have an important role selecting animal traits and provides useful information on how to build more biodiversity-friendly cities.

The episodic resurgence of highly pathogenic avian influenza H5 virus.

Highly pathogenic avian influenza (HPAI) H5N1 activity has intensified globally since 2021, increasingly causing mass mortality in wild birds and poultry and incidental infections in mammals1-3. However, the ecological and virological properties that underscore future mitigation strategies still remain unclear. Using epidemiological, spatial and genomic approaches, we demonstrate changes in the origins of resurgent HPAI H5 and reveal significant shifts in virus ecology and evolution. Outbreak data show key resurgent events in 2016-2017 and 2020-2021, contributing to the emergence and panzootic spread of H5N1 in 2021-2022. Genomic analysis reveals that the 2016-2017 epizootics originated in Asia, where HPAI H5 reservoirs are endemic. In 2020-2021, 2.3.4.4b H5N8 viruses emerged in African poultry, featuring mutations altering HA structure and receptor binding. In 2021-2022, a new H5N1 virus evolved through reassortment in wild birds in Europe, undergoing further reassortment with low-pathogenic avian influenza in wild and domestic birds during global dissemination. These results highlight a shift in the HPAI H5 epicentre beyond Asia and indicate that increasing persistence of HPAI H5 in wild birds is facilitating geographic and host range expansion, accelerating dispersion velocity and increasing reassortment potential. As earlier outbreaks of H5N1 and H5N8 were caused by more stable genomic constellations, these recent changes reflect adaptation across the domestic-bird-wild-bird interface. Elimination strategies in domestic birds therefore remain a high priority to limit future epizootics.

AN EVALUATION OF SYLVIIDAE SPECIES IN THE TRANSPORT OF POLLEN: A CASE STUDY OF İSTANBUL

During pollination, pollen grains can be transported to distant distances by various factors such as wind, insects, birds, and reptiles. While there have been numerous studies on pollen transportation by birds in Europe, there is a lack of research concerning the pollen grains transported by bird species in Turkey. In this context, we explored the role of Sylviidae species in pollination and identified the plant species whose pollen is transported by these birds. The temporary ringing station located at İstanbul University-Cerrahpaşa Avcılar Campus was selected as the study area and a total of 35 feather samples were collected from individuals of Sylvia atricapilla (Linnaeus), S. borin (Boddaert), Curruca communis (Latham), C. cantillans (Pallas), C. curruca (Linnaeus) and C. melanocephala (Gmelin) captured at station. Based on the analysis of pollen found on the feather samples, it was determined that the pollen found in 18 feather samples of Curruca curruca and Sylvia atricapilla provided pollen belonging to Pinus (Pine), Aesculus (Horse chestnut), Prunus (Plum), Spiraea (Spiraea), Quercus (Oak), Fraxinus (Ash) and Melia (Bead tree). The most commonly determined pollen on C. curruca were of Prunus (63.5%), while the least common were of Aesculus (0.9%). The most commonly determined pollen on S. atricapilla were of Melia (48.1%), while the least common were of Crocus (0.2%). The presence of pollen among the food sources of Sylviidae species during spring, the synchronization between the arrival time of birds in the study area during migration and the pollination period of the determined plant taxa, and the prevalence of these taxa in the study area suggest that S. atricapilla and C. curruca may be contributing to pollination.

Epidemiology of emerging viruses

Newly emerging viruses such as West Nile virus, coronaviruses (SARS-CoV, MERS-CoV and SARS-CoV-2), H1N1pdm09, Ebola virus, Zika virus and Monkeypox virus are of global concern as they have caused serious public threats. Among the several outbreaks caused by emerging viruses, the worst pandemic was caused by SARS-CoV-2. The Omicron variant is the dominant variant among SARS-CoV-2 variants and the lineages BA.5.2 and BF.7 were responsible for the recent Chinese surge. Genome sequencing studies of MERS-CoV revealed that it is evolving with mutations in spike protein. West Nile virus is still circulating among birds in Europe, Mediterranean and North American countries. Several West Nile virus cases were reported from Europe and neighboring countries in 2022. The 2009 H1N1 pandemic caused by H1N1pdm09 is a variant strain evolved from H1N1 in swine. Though it caused more than 60 million infections, the mortality rate was 0.02%. Staggeringly, severe infections occurred among obese people. Zika virus has jumped borders from Africa and is adapting to become endemic in new geographical habitats such as South East Asia, India and Europe due to uncontrolled breeding of Aedes mosquitoes. Recently, autochthonous vector borne transmission of Zika virus was documented in France. During the Monkeypox pandemic, the virus demonstrated a lower infectious and mortality rate in individuals vaccinated for smallpox. A lower virulence of Monkeypox virus compared to smallpox was due to mutations in the ORF regions. Surveillance efforts and measures must be incessantly undertaken to prevent future outbreaks caused by emerging viruses.

Sarcocystis spp. Macrocysts Infection in Wildfowl Species in Eastern Baltic Region: Trends in Prevalence in 2011-2022.

Wildfowl meat infected with S. rileyi macrocysts is not suitable for human consumption. Ducks are among the main game birds in Europe, and S. rileyi infections cause significant economic losses. In 2011-2022, a total of 2649 anseriforms collected in Lithuania and 619 Mallards (Anas platyrhynchos) hunted in the Kaliningrad region of Russia, Belarus, and Latvia were tested for macrocysts. In Lithuania, macrocysts were detected in 206 of 2362 Mallards (8.7%) and in two of 88 (2.3%) Eurasian Teals (Anas crecca). The prevalence of macrocysts in the other three countries, Belarus (5.9%), Russia (5.0%), and Latvia (3.1%), was similar. For species identification, macrocysts isolated from 37 Mallards (21 from Lithuania, 8 from Russia, 6 from Belarus, and 2 from Latvia) were subjected to sequencing of the ITS1 region. Based on DNA analysis, S. rileyi was confirmed in all tested birds. By comparing the infection rates of macrocysts in Mallards in Lithuania, significant differences were observed in different years (p = 0.036), and a significantly higher prevalence of infection was established in November-December than in September-October (p = 0.028). Given the amount of data per decade on the prevalence of S. rileyi, awareness of infection needs to be increased.

Trends in the global trade of live CITES-listed raptors: Trade volumes, spatiotemporal dynamics and conservation implications

The global legal wildlife trade is worth US$4–20 billion to the world's economy every year. Raptors frequently enter the wildlife trade for use as display animals, by falconers or hobbyists for sport and recreation. Using data from the Convention on International Trade in Endangered Species of Wild Fauna and Flora's (CITES) Trade Database, we examined trends in the global, legal commercial trade of CITES-listed raptors between 1975 and 2020. Overall 272 species were traded, totalling 188,149 traded individuals, which increased over time. Hybrid Falcons (N = 50,366) were most commonly traded, comprising more than a third of the global diurnal CITES-listed raptor trade, followed by Gyrfalcons (Falco rusticolus; N = 30,510), Saker Falcons (F. cherrug; N = 21,679), Peregrine Falcons (F. peregrinus; N = 13,390) and Northern White-faced Owls (Ptilopsis leucotis; N = 6725). More than half of wild-caught diurnal raptors were classified as globally threatened. The United Kingdom was the largest exporter of live raptors and the United Arab Emirates was the largest importer. Countries with higher GDPs (US$) imported more raptors than those with smaller GDPs. Larger-bodied diurnal species were traded more relative to smaller-bodied conspecifics. Following the introduction of the European Union's Wild Bird Trade Ban in 2005, the number of traded wild-caught raptors declined. Despite its limitations, the CITES Trade Database provides an important baseline of the global legal trade of live raptors. However, better understanding of illegal wildlife trade networks and smuggling routes, both on-the-ground and online, are essential for future conservation efforts.

Local colonisations and extinctions of European birds are poorly explained by changes in climate suitability

Climate change has been associated with both latitudinal and elevational shifts in species’ ranges. The extent, however, to which climate change has driven recent range shifts alongside other putative drivers remains uncertain. Here, we use the changing distributions of 378 European breeding bird species over 30 years to explore the putative drivers of recent range dynamics, considering the effects of climate, land cover, other environmental variables, and species’ traits on the probability of local colonisation and extinction. On average, species shifted their ranges by 2.4 km/year. These shifts, however, were significantly different from expectations due to changing climate and land cover. We found that local colonisation and extinction events were influenced primarily by initial climate conditions and by species’ range traits. By contrast, changes in climate suitability over the period were less important. This highlights the limitations of using only climate and land cover when projecting future changes in species’ ranges and emphasises the need for integrative, multi-predictor approaches for more robust forecasting.

Generation time and seasonal migration explain variation in spatial population synchrony across European bird species.

Spatial population synchrony is common among populations of the same species and is an important predictor of extinction risk. Despite the potential consequences for metapopulation persistence, we still largely lack understanding of what makes one species more likely to be synchronized than another given the same environmental conditions. Generally, environmental conditions in a shared environment or a species' sensitivity to the environment can explain the extent of synchrony. Populations that are closer together experience more similar fluctuations in their environments than those populations that are further apart and are therefore more synchronized. The relative importance of environmental and demographic stochasticity for population dynamics is strongly linked to species' life-history traits, such as pace of life, which may impact population synchrony. For populations that migrate, there may be multiple environmental conditions at different locations driving synchrony. However, the importance of life history and migration tactics in determining patterns of spatial population synchrony have rarely been explored empirically. We therefore hypothesize that increasing generation time, a proxy for pace of life, would decrease spatial population synchrony and that migrants would be less synchronized than resident species. We used population abundance data on breeding birds from four countries to investigate patterns of spatial population synchrony in growth rate and abundance. We calculated the mean spatial population synchrony between log-transformed population growth rates or log-transformed abundances for each species and country separately. We investigated differences in synchrony across generation times in resident (n = 67), short-distance migrant (n = 86) and long-distance migrant (n = 39) bird species. Species with shorter generation times were more synchronized than species with longer generation times. Short-distance migrants were more synchronized than long-distance migrants and resident birds. Our results provide novel empirical links between spatial population synchrony and species traits known to be of key importance for population dynamics, generation time and migration tactics. We show how these different mechanisms can be combined to understand species-specific causes of spatial population synchrony. Understanding these specific drivers of spatial population synchrony is important in the face of increasingly severe threats to biodiversity and could be key for successful future conservation outcomes.

Highly pathogenic avian influenza A (H5N1) virus infections in wild carnivores connected to mass mortalities of pheasants in Finland.

Highly pathogenic avian influenza (HPAI) has caused widespread mortality in both wild and domestic birds in Europe during 2020-2022. Virus types H5N8 and H5N1 have dominated the epidemic. Isolated spill-over infections in mammals started to emerge as the epidemic continued. In autumn 2021, HPAI H5N1 caused a series of mass mortality events in farmed and released pheasants (Phasianus colchicus) in a restricted area in southern Finland. Later, in the same area, an otter (Lutra lutra), two red foxes (Vulpes vulpes) and a lynx (Lynx lynx) were found moribund or dead and infected with H5N1 HPAI virus. Phylogenetically, H5N1 strains from pheasants and mammals clustered together. Molecular analyses of the four mammalian virus strains revealed mutations in the PB2 gene segment (PB2-E627K and PB2-D701N) that are known to facilitate viral replication in mammals. This study revealed that avian influenza cases in mammals were spatially and temporally connected with avian mass mortalities suggesting increased infection pressure from birds to mammals.

Comparative Analysis of the Exo-Erythrocytic Development of Five Lineages of Haemoproteus majoris, a Common Haemosporidian Parasite of European Passeriform Birds.

Haemoproteus parasites (Apicomplexa, Haemosporida) are widespread pathogens of birds, with a rich genetic (about 1900 lineages) and morphospecies (178 species) diversity. Nonetheless, their life cycles are poorly understood. The exo-erythrocytic stages of three Haemoproteus majoris (widespread generalist parasite) lineages have been previously reported, each in a different bird species. We aimed to further study and compare the development of five H. majoris lineages-hCCF5, hCWT4, hPARUS1, hPHSIB1, and hWW2-in a wider selection of natural avian hosts. A total of 42 individuals belonging to 14 bird species were sampled. Morphospecies and parasitemia were determined by microscopy of blood films, lineages by DNA-barcoding a 478 bp section of the cytochrome b gene, and exo-erythrocytic stages by histology and chromogenic in situ hybridization. The lineage hCWT4 was morphologically characterized as H. majoris for the first time. All lineage infections exclusively featured megalomeronts. The exo-erythrocytic stages found in all examined bird species were similar, particularly for the lineages hCCF5, hPARUS1, and hPHSIB1. Megalomeronts of the lineages hWW2 and hCWT4 were more similar to each other than to the former three lineages. The kidneys and gizzard were most often affected, followed by lungs and intestines; the site of development showed variation depending on the lineage.