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Background Avian malaria parasites and related haemosporidians are diverse and widespread vector-borne parasites that circulate within complex host-vector networks. Despite their ecological importance, their diversity and circulation within mosquito communities remain poorly understood in temperate North America. The Midwestern United States, located along major North American migratory flyways, supports diverse avian habitats and mosquito assemblages, providing an ideal setting to investigate these interactions. Here, we characterized the occurrence, phylogenetic diversity, and mosquito-lineage associations of avian haemosporidians in mosquitoes. Methods A total of 233 pools comprising 6170 unfed female mosquitoes from seven species were collected from 38 protected natural areas in northern, central, and southern Illinois. Samples were screened for avian haemosporidian DNA by nested PCR targeting the mitochondrial cytochrome b gene. Positive samples were analyzed by multiplex PCR and sequencing to identify parasite lineages and assess potential mixed infections. Phylogenetic relationships were reconstructed, and mosquito-lineage associations were visualized using a bipartite network. Infection rates were estimated using maximum likelihood estimation (MLE) and minimum infection rate (MIR), and factors associated with infection prevalence in pooled samples were evaluated using pooled-binomial regression. Results Of 233 pools, 44 (18.9%) were positive for avian haemosporidians. Eleven lineages were identified, including nine Plasmodium and two Haemoproteus lineages; one Plasmodium lineage was novel. pTUMIG03 ( Plasmodium unalis ) was the most frequently detected lineage, whereas pSYAT05 ( P. vaughani ) showed the broadest mosquito distribution. Culex restuans harbored the highest lineage diversity and infection rate among the mosquito species examined. Mosquito species was a significant predictor of infection prevalence estimated from pooled samples, and Cx. restuans was most strongly associated with avian Plasmodium detection. Additionally, a deer-associated non-avian Plasmodium lineage was incidentally detected in two pools. Conclusions Mosquito populations in Illinois harbor a diverse assemblage of avian haemosporidians, with heterogeneous lineage distributions across mosquito taxa. Culex restuans emerged as the species most strongly associated with avian Plasmodium prevalence, suggesting an important role in local enzootic circulation. These findings expand current knowledge of avian haemosporidian diversity in North American mosquito communities and provide a basis for future studies integrating mosquito surveillance, avian host sampling, and vector competence experiments.

Birds possess the most diverse assemblage of haemosporidian parasites, although the true diversity is unknown due to high genetic diversity and insufficient sampling across all avian clades. Waterfowl (Order Anseriformes) are an ideal group to discover hidden parasite diversity and examine the role of host ecology in parasite transmission. Waterfowl contain 2 distinct feeding guilds, dabbling and diving, which differ in niche utilization that likely alters vector encounter rates and haemosporidian parasite risk. To determine the role of feeding guild in haemosporidian parasitism we analysed 223 blood samples collected by hunters from the upper Midwest of the United States from 2017 to 2019. Fifty-four individuals were infected by haemosporidian parasites (24·2% prevalence). Infection prevalence differed significantly between dabbling (34·9%, n = 109) and diving (14·0%, n = 114) ducks. Feeding guild was the only host trait that could predict haemosporidian infection risk, with a significantly higher risk in dabbling ducks. Twenty-four haemosporidian lineages were identified, with 9 identified for the first time. Thirteen lineages were found only in dabbling ducks, 5 only in diving ducks and 6 in both feeding guilds. Community analysis showed that each feeding guild harboured a unique parasite community. There was no phylogenetic signal of feeding guild within a phylogenetic reconstruction of North American waterfowl haemosporidian lineages. Our results demonstrate that waterfowl contain a diverse and distinct community of haemosporidian parasites. The unique composition of each feeding guild determines not only haemosporidian infection risk but also community structure. This is the first report of such an impact for waterfowl feeding guilds.

Haemoproteus bobricklefsi sp. nov. (Haemosporida, Haemoproteidae) was found in the dunnock Prunella modularis and represents the first blood parasite described in accentor birds of the Prunellidae. The description is based on the morphology of blood stages and includes information about a barcoding segment of the mitochondrial cytochrome b gene (lineage hDUNNO01) and the full mitochondrial genome, which can be used for identification and diagnosis of this infection. The new parasite can be readily distinguished from described species of haemoproteids parasitizing passeriform birds due to markedly variable position of nuclei in advanced and fully grown macrogametocytes. Illustrations of blood stages of the new species are given, and phylogenetic analyses based on partial mitochondrial cytochrome b gene sequences and the full mitochondrial genome identified the closely related lineages. DNA haplotype networks showed that transmission occurs in Europe and North America. This parasite was found in the dunnock in Europe and several species of the Passerellidae in North America. It is probably of Holarctic distribution, with the highest reported prevalence in the UK. The parasite distribution seems to be geographically patchy, with preference for areas of relatively cool climates. Phylogenetic analysis suggests that H. bobricklefsi sp. nov. belongs to the Parahaemoproteus subgenus and is probably transmitted by biting midges belonging to Culicoides (Ceratopogonidae). The available data on molecular occurrence indicate that this pathogen is prone to abortive development, so worth attention in regard of consequences for bird health.

Biodiversity varies worldwide and is influenced by multiple factors, such as environmental stability and past historical events (e.g. Panama Isthmus). At the same time, organisms with unique life histories (e.g. parasites) are subject to unique selective pressures that structure their diversity patterns. Parasites represent one of the most successful life strategies, impacting, directly and indirectly, ecosystems by cascading effects on host fitness and survival. Here, I focused on a highly diverse, prevalent and cosmopolitan group of parasites (avian haemosporidians) to investigate the main drivers (e.g. host and environmental features) of regional parasite diversity on a global scale. To do so, I compiled data from 4 global datasets on (i) avian haemosporidian (malaria and malaria-like) parasites, (ii) bird species diversity, (iii) avian functional traits and (iv) climate data. Then, using generalized least square models, I evaluated the effect of host and environmental features on haemosporidian diversity. I found that haemosporidian diversity mirrors host regional diversity and that higher host body mass increases haemosporidian diversity. On the other hand, climatic conditions had no effect on haemosporidian diversity in any model. When evaluating Leucocytozoon parasites separately, I found parasite diversity was boosted by a higher proportion of migratory hosts. In conclusion, I demonstrated that haemosporidian parasite diversity is intrinsically associated with their hosts’ diversity and body mass.

Black flies (Diptera: Simuliidae) are significant pests and vectors transmitting pathogens to humans and other animals. However, knowledge of species diversity as well as their role as pests and vectors in Laos is limited, despite the country having abundant suitable lotic habitats, that is, running water. In this study, we collected wild adult black fly specimens from Laos. Both morphology and genetic data supported that they were Simulium khelangense Takaoka, Srisuka and Saeung, and therefore, a new distribution record for this species. Screening of the blood protozoa from 106 females detected Leucocytozoon (n = 7) and Trypanosoma (n = 9). Identifications of Leucocytozoon based on cytochrome b sequences in MalAvi database indicated that 5 specimens had identical sequences to those of 4 lineages (GALLUS17 [n = 2], GALLUS35 [n = 1], GALLUS37 [n = 1], and GALLUS41 [n = 1]) of unidentified species (i.e., Leucocytozoon sp.). The remaining 2 had no identical sequences but were very similar (99%) to GALLUS34 and GALLUS41 lineages. Identifications of Trypanosoma based on the small subunit rRNA indicated that all were T. avium. The results presented here further support the role of S. khelangense as a potential vector of Leucocytozoon and Trypanosoma as well as being a possible pest of domestic avian species.

Migratory animals likely play an important role in the geographic spread of parasites. In fact, a common assumption is that parasites are potentially transmitted by migratory animals at temporary stopover sites along migratory routes, yet very few studies have assessed whether transmission at stopover sites can or does occur. We investigated the potential for a group of vector-transmitted parasites, the avian haemosporidians, to be transmitted during migratory stopover periods at Rushton Woods Preserve in Pennsylvania, USA. Using an analysis of 1454 sampled avian hosts, we found that while a core group of abundant haemosporidians was shared between local breeding birds and passing migrants, the parasite community of migratory birds at Rushton was distinct from that of local breeding birds and showed similarity to a previously sampled boreal forest haemosporidian community. Haemosporidians that were unique to passing migratory birds were associated with sampling sites in North America with cooler summer temperatures than haemosporidians that are transmitted at Rushton, suggesting that the transmission of these parasites may be restricted to high-latitude regions outside of our temperate stopover site. We also found that the abundance of mosquitoes in our study region is offset from that of migratory bird abundance during avian migratory periods, with the peak period of bird migration occurring during periods of low mosquito activity. Collectively, these findings suggest that although abundant haemosporidians are possibly transmitted between local and passing migratory birds, a combination of biotic and abiotic factors may constrain haemosporidian transmission during avian stopover at our study site.

Background: The order Accipitriformes comprises the largest group of birds of prey with 260 species in four families. So far, 21 haemosporidian parasite species have been described from or reported to occur in accipitriform birds. Only five of these parasite species have been characterized molecular genetically. The first part of this study involved molecular genetic screening of accipitriform raptors from Austria and Bosnia-Herzegovina and the first chromogenic in situ hybridization approach targeting parasites in this host group. The aim of the second part of this study was to summarize the CytB sequence data of haemosporidian parasites from accipitriform raptors and to visualize the geographic and host distribution of the lineages. Methods: Blood and tissue samples of 183 accipitriform raptors from Austria and Bosnia-Herzegovina were screened for Plasmodium, Haemoproteus and Leucocytozoon parasites by nested PCR, and tissue samples of 23 PCR-positive birds were subjected to chromogenic in situ hybridization using genus-specific probes targeting the parasites’ 18S rRNAs. All published CytB sequence data from accipitriform raptors were analysed, phylogenetic trees were calculated, and DNA haplotype network analyses were performed with sequences from clades featuring multiple lineages detected in this host group. Results: Of the 183 raptors from Austria and Bosnia-Herzegovina screened by PCR and sequencing, 80 individuals (44%) were infected with haemosporidian parasites. Among the 39 CytB lineages detected, 18 were found for the first time in the present study. The chromogenic in situ hybridization revealed exo-erythrocytic tissue stages of Leucocytozoon parasites belonging to the Leucocytozoon toddi species group in the kidneys of 14 infected birds. The total number of CytB lineages recorded in accipitriform birds worldwide was 57 for Leucocytozoon, 25 for Plasmodium, and 21 for Haemoproteus. Conclusion: The analysis of the DNA haplotype networks allowed identifying numerous distinct groups of lineages, which have not yet been linked to morphospecies, and many of them likely belong to yet undescribed parasite species. Tissue stages of Leucocytozoon parasites developing in accipitriform raptors were discovered and described. The majority of Leucocytozoon and Haemoproteus lineages are specific to this host group, but most Plasmodium lineages were found in birds of other orders. This might indicate local transmission from birds kept at the same facilities (raptor rescue centres and zoos), likely resulting in abortive infections. To clarify the taxonomic and systematic problems, combined morphological and molecular genetic analyses on a wider range of accipitriform host species are needed.

Blood parasites from the order Haemosporida infect many vertebrates and cause malaria-like diseases. In this study, a haemosporidian infection was detected in a sick grey crowned crane imported into China using a combination of morphological and molecular approaches. Blood samples were collected from the jugular vein and processed for morphological identification of infective parasites using stained blood smears and microscopy. No merogony occurs in the blood cells, and sporadic pigment granules were observed. Nested-PCR assays were employed for a molecular examination, which indicated that the cytb gene of this parasite had 94.1–94.9% identity to Haemoproteus antigonis . Subsequently, its mitochondrial genome structure was determined by high-throughput sequencing using the DNBSEQ-T7 platform. The determined structure was confirmed by the Sanger sequencing using amplicons. The mitochondrial genome obtained for this parasite exhibited a low CG content (32.0%) and possessed three protein-coding genes, encoding 1068 amino acids, which constituted 53.7% of the genome. Phylogenetic analysis indicated that this parasite clustered with Haemoproteus sp . is detected in grey crowned cranes from Africa. This parasite was likely acquired during importation of this animal; thus, strict quarantine of imported ornamental animals is required to prevent the entry of new pathogens.

Background: Haemosporidians (Haemosporida, Apicomplexa), which include malaria parasites, are found in nearly all terrestrial ecosystems. Avian haemosporidians have been extensively studied; however, there is limited information on parasites in owls (order Strigiformes). Here, haemosporidians infecting North American owls were characterized using an integrative methodology. Methods: Taking advantage of injured/dead owls from rehabilitation centres in the central and northeastern USA, 53 individuals of Bubo virginianus, Strix varia, Megascops asio, Bubo scandiacus, Aegolius acadicus, and Tyto furcata were screened using polymerase chain reaction for all samples and microscopy for those with available blood smears. Parasite mitochondrial genomes were obtained using a long-read sequencing method (PacBio HiFi), which efficiently detects multiple infections in a single host. The relationships between parasite lineages were estimated using phylogenetic and haplotype network methods. Results: In total, 21 individuals from three species were positive by PCR: B. virginianus (14/17, 82.4%), M. asio (3/5, 60%), and S. varia (4/8, 50%). Two Plasmodium, three Haemoproteus, and four Leucocytozoon lineages were identified infecting these hosts, with one Haemoproteus and one Leucocytozoon being new to science. All positive individuals were infected with Haemoproteus parasites, and two B. virginianus had a mixed infection with Leucocytozoon and Haemoproteus species. The hSTVAR01 cytochrome b (cytb) lineage common in North America is linked for the first time to Haemoproteus syrnii. Haemoproteus syrnii was found in all 14 positive B. virginianus and two S. varia. Notably, all the cytb lineages from previously identified H. syrnii, based on erythrocytic stages, were not monophyletic, indicating the existence of an undescribed species. The pPADOM11 cytb lineage was recognized as an allele of Plasmodium elongatum. Conclusion: Long reads enabled the detection of mixed/co-infections. The link between genetic data and morphospecies was established in two cases. Several Leucocytozoon clades were observed; however, only one morphospecies, Leucocytozoo danilewskyi, has been described in owls. Thus, there is a need for a detailed analysis of blood stages to determine whether different owl Haemoproteus and Leucocytozoon parasites exhibit morphological differences or represent cryptic species. Overall, this study underscores the importance of high-quality molecular data in characterizing the biodiversity of haemosporidian parasites.

Haemosporida are vector-borne parasitic protozoa known to be present in birds of most avian orders. However, despite their perceived diversity using DNA barcode approaches, describing and delimiting species is challenging, particularly for those parasites found in non-passerine birds. In this study, we describe Haemoproteus trarotraro n. sp., a species found in two Falconiform hosts, the Crested Caracara ( Caracara plancus plancus , type host) and the Yellow-headed Caracara ( Daptrius chimachima chimachima ), both sampled in Brazil at a wildlife rehabilitation center using microscopy and molecular tools. Haemoproteus trarotraro n. sp. is distinguished from the two other haemoproteid species described in Falconiformes, H. brachiatus and H. tinnunculi , by the absence of gametocytes that fully encircle the host-cell nucleus, and by the presence of numerous small vacuoles scattered throughout the cytoplasm of macrogametocytes. Both the partial cytb gene and the mtDNA genome for this new species are reported. The sequencing of the cytb barcode fragment revealed that H. trarotraro n. sp. reported here corresponds to a Haemoproteus sp. haplotype (GenBank Accession (AF465594) lineage POLPLA01 in Malavi) previously reported from Caracara plancus cheriway in Florida, USA. Although it diverges by only  2% at the cytb level from H. tinnunculi and H. brachiatus , H. trarotraro n. sp. is not a sister lineage to these taxa. Instead, phylogenetic analyses place it within a distinct but closely related, well-supported clade comprising lineages infecting American Kestrels ( Falco sparverius ). This study contributes, through an integrative taxonomic approach, to the ongoing discussion about species delimitation within the order Haemosporida.