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Gediminas Valkiūnas - One of the best experts on this subject based on the ideXlab platform.
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Penguins are competent hosts of Haemoproteus parasites: the first detection of gametocytes, with molecular characterization of Haemoproteus larae
Parasites & vectors, 2020Co-Authors: Mizue Inumaru, Yukita Sato, Shiori Aratani, Misa Shimizu, Mineka Yamamoto, Koichi Murata, Gediminas ValkiūnasAbstract:Background The majority of penguins (Sphenisciformes) have evolved in areas with weak or absent transmission of haemosporidian parasites and are usually naive to avian haemosporidian infections. Plasmodium parasites are transmitted by mosquitoes, and lethal avian malaria has been often reported in captive penguins in many countries. The related haemosporidian parasites belonging to Haemoproteus and Leucocytozoon have also been detected in penguins but less often than Plasmodium infections. The majority of Haemoproteus infection reports in penguins are based solely on PCR-based diagnostics. It remains unclear if haemoproteids can complete their life-cycle and produce infective stages (gametocytes) in penguins or whether these infections are abortive in penguins, and thus dead ends for transmission. In other words, it remains unknown if penguins are competent hosts for Haemoproteus parasites, which cause disease in non-adapted birds. Methods Two captive African penguins (Spheniscus demersus) and two Magellanic penguins (S. magellanicus) were found to be positive for Haemoproteus infection in two open-air aquariums in Japan, and the parasites were investigated using both PCR-based testing and microscopical examination of blood films. Samples from a black-tailed gull (Larus crassirostris) and previously tested gulls were used for comparison. Results The lineage hSPMAG12 was detected, and gametocytes of Haemoproteus sp. were seen in the examined penguins and gull. Observed gametocytes were indistinguishable from those of Haemoproteus larae, which naturally parasitize birds of the genus Larus (Laridae). The detected sequence information and Bayesian phylogenetic analysis supported this conclusion. Additionally, morphologically similar gametocytes and closely related DNA sequences were also found in other gull species in Japan. Phylogenetic analysis based on partial cytb sequences placed the lineage hSPMAG12 of H. larae within the clade of avian haemoproteids which belong to the subgenus ParaHaemoproteus, indicating that Culicoides biting midges likely transmit the parasites between penguins and gulls. Conclusions This study shows that some species of Haemoproteus parasites complete their development and produce gametocytes in penguins, which may be source of infection for biting midges transmitting haemoproteosis. To prevent haemosporidiosis in zoos, we call for control not only of mosquitoes, but also biting midges.
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Sporogony of four Haemoproteus species (Haemosporida: Haemoproteidae), with report of in vitro ookinetes of Haemoproteus hirundinis: phylogenetic inference indicates patterns of haemosporidian parasite ookinete development.
Parasites & vectors, 2019Co-Authors: Carolina Romeiro Fernandes Chagas, Rasa Bernotienė, Tatjana A. Iezhova, Dovilė Bukauskaitė, Mikas Ilgūnas, Gediminas ValkiūnasAbstract:Haemoproteus (ParaHaemoproteus) species (Haemoproteidae) are widespread blood parasites that can cause disease in birds, but information about their vector species, sporogonic development and transmission remain fragmentary. This study aimed to investigate the complete sporogonic development of four Haemoproteus species in Culicoides nubeculosus and to test if phylogenies based on the cytochrome b gene (cytb) reflect patterns of ookinete development in haemosporidian parasites. Additionally, one cytb lineage of Haemoproteus was identified to the species level and the in vitro gametogenesis and ookinete development of Haemoproteus hirundinis was characterised. Laboratory-reared C. nubeculosus were exposed by allowing them to take blood meals on naturally infected birds harbouring single infections of Haemoproteus belopolskyi (cytb lineage hHIICT1), Haemoproteus hirundinis (hDELURB2), Haemoproteus nucleocondensus (hGRW01) and Haemoproteus lanii (hRB1). Infected insects were dissected at intervals in order to detect sporogonic stages. In vitro exflagellation, gametogenesis and ookinete development of H. hirundinis were also investigated. Microscopic examination and PCR-based methods were used to confirm species identity. Bayesian phylogenetic inference was applied to study the relationships among Haemoproteus lineages. All studied parasites completed sporogony in C. nubeculosus. Ookinetes and sporozoites were found and described. Development of H. hirundinis ookinetes was similar both in vivo and in vitro. Developing ookinetes of this parasite possess long outgrowths, which extend longitudinally and produce the apical end of the ookinetes. A large group of closely related Haemoproteus species with a similar mode of ookinete development was determined. Bayesian analysis indicates that this character has phylogenetic value. The species identity of cytb lineage hDELURB2 was determined: it belongs to H. hirundinis. Culicoides nubeculosus is susceptible to and is a likely natural vector of numerous species of Haemoproteus parasites, thus worth attention in haemoproteosis epidemiology research. Data about in vitro development of haemoproteids provide valuable information about the rate of ookinete maturation and are recommended to use as helpful step during vector studies of haemosporidian parasites, particularly because they guide proper dissection interval of infected insects for ookinete detection during in vivo experiments. Additionally, in vitro studies readily identified patterns of morphological ookinete transformations, the characters of which are of phylogenetic value in haemosporidian parasites.
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Molecular characterization of six widespread avian haemoproteids, with description of three new Haemoproteus species.
Acta tropica, 2019Co-Authors: Gediminas Valkiūnas, Rasa Bernotienė, Dovilė Bukauskaitė, Mikas Ilgūnas, Carolina Romeiro Fernandes Chagas, Tanja Himmel, Josef Harl, Herbert Weissenböck, Tatjana A. IezhovaAbstract:Abstract Species of Haemoproteus (Haemosporida, Haemoproteidae) are widespread and often prevalent blood parasites of birds all over the word. They are particularly diverse in tropical countries. Due to limited knowledge of life cycles, these pathogens usually have been considered relatively benign and were neglected in veterinary medicine and bird management. However, recent molecular studies provided evidence that Haemoproteus parasites might cause severe diseases if they infect non–adapted (wrong) avian hosts due to marked damage of organs by exo-erythrocytic stages (megalomeronts). Additionally, high Haemoproteus infections are lethal to blood-sucking insects. Molecular markers are essential for reliable detection and species identification both at tissue stages in vertebrates and sporogonic stages in arthropods however, remain insufficiently developed for wildlife haemosporidian parasites. This study combined PCR-based and microscopic approaches and reported cytochrome b gene (cytb) and apicoplast gene (clpc) markers for characterization of six widespread species of haemoproteids parasitizing common birds wintering in tropics and subtropics of the Old World. Three new Haemoproteus species were described using morphological and molecular markers. Molecular characterization of haemoproteids parasitizing falcons was developed. Morphological and phylogenetic characterization of Haemoproteus tinnunculi (cytb lineage hFALSUB01), H. brachiatus (hLK03), H. parabelopolskyi (hSYAT1), H. homogeneae n. sp. (hSYAT16), H. homopicae n. sp. (hGAGLA07) and H. homominutus n. sp. (hCUKI1) was performed and provides clues for infections diagnostics. This study adds three species to the group of morphologically readily distinct Haemoproteus parasites, which differ in few base pairs (
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Molecular prevalence and phylogenetic relationship of Haemoproteus and Plasmodium parasites of owls in Thailand: Data from a rehabilitation centre.
International journal for parasitology. Parasites and wildlife, 2019Co-Authors: Pornchai Pornpanom, Gediminas Valkiūnas, Chaiyan Kasorndorkbua, Preeda Lertwatcharasarakul, Carolina Romeiro Fernandes Chagas, Chaleow SalakijAbstract:Owls are nocturnal raptors that are prevalently infected with haemosporidian parasites wordwide. These birds were commonly submitted to the Kasetsart University Raptor Rehabilitation Unit, Kasetsart University, Thailand and were examined using PCR-based methods for the presence of haemosporidian infections of by the genera Plasmodium and Haemoproteus. Blood samples from 167 individual owls belonging to 12 species common in Thailand were collected between September 2012 and February 2018. The overall prevalence of haemosporidians was 34.1%, with Haemoproteus infections (25.1%) being more prevalent than Plasmodium infections (9.0%). The prevalence of both Haemoproteus and Plasmodium parasites was similar in all seasons of the year. Molecular characterization revealed 17 new haemosporidian parasite lineages (11 Haemoproteus and six Plasmodium), with genetic variation among partial cytochrome b sequences ranging from 0.0% to 3.6% in Haemoproteus lineages and 0.2%-8.8% in Plasmodium lineages. Phylogenetic analysis showed that all Haemoproteus lineages detected in owls appeared in one well-supported clade together with other parasites belonging to the ParaHaemoproteus subgenus, indicating their close evolutionary relationship and common transmission modality by Culicoides biting midges. This study showes the existence of prominent non-described haemosporidian parasite diversity in Thai owls and provides baseline molecular information for further research on the genetic diversity of owl haemosporidian parasites. New DNA sequence information can be used for the diagnosis of owl infections, which have been often reported during rehabilitation planning.
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Delineation of the Genera Haemoproteus and Plasmodium Using RNA-Seq and Multi-gene Phylogenetics
Journal of molecular evolution, 2018Co-Authors: Jasper Toscani Field, Staffan Bensch, Gediminas Valkiūnas, Josh Weinberg, Nubia E. Matta, Ravinder N. M. SehgalAbstract:Members of the order Haemosporida are protist parasites that infect mammals, reptiles and birds. This group includes the causal agents of malaria, Plasmodium parasites, the genera Leucocytozoon and Fallisia, as well as the species rich genus Haemoproteus with its two subgenera Haemoproteus and ParaHaemoproteus. Some species of Haemoproteus cause severe disease in avian hosts, and these parasites display high levels of diversity worldwide. This diversity emphasizes the need for accurate evolutionary information. Most molecular studies of wildlife haemosporidians use a bar coding approach by sequencing a fragment of the mitochondrial cytochrome b gene. This method is efficient at differentiating parasite lineages but insufficient for accurate phylogenetic inferences in highly diverse taxa such as haemosporidians. Recent studies have utilized multiple mitochondrial genes (cyt b, cox1 and cox3), sometimes combined with a few apicoplast and nuclear genes. These studies have been highly successful with one notable exception: the evolutionary relationships of the genus Haemoproteus remain unresolved. Here we describe the transcriptome of Haemoproteus columbae and investigate its phylogenetic position recovered from a multi-gene dataset (600 genes). This genomic approach restricts the taxon sampling to 18 species of apicomplexan parasites. We employed Bayesian inference and maximum likelihood methods of phylogenetic analyses and found H. columbae and a representative from the subgenus ParaHaemoproteus to be sister taxa. This result strengthens the hypothesis of genus Haemoproteus being monophyletic; however, resolving this question will require sequences of orthologs from, in particular, representatives of Leucocytozoon species.
Tatjana A. Iezhova - One of the best experts on this subject based on the ideXlab platform.
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Sporogony of four Haemoproteus species (Haemosporida: Haemoproteidae), with report of in vitro ookinetes of Haemoproteus hirundinis: phylogenetic inference indicates patterns of haemosporidian parasite ookinete development.
Parasites & vectors, 2019Co-Authors: Carolina Romeiro Fernandes Chagas, Rasa Bernotienė, Tatjana A. Iezhova, Dovilė Bukauskaitė, Mikas Ilgūnas, Gediminas ValkiūnasAbstract:Haemoproteus (ParaHaemoproteus) species (Haemoproteidae) are widespread blood parasites that can cause disease in birds, but information about their vector species, sporogonic development and transmission remain fragmentary. This study aimed to investigate the complete sporogonic development of four Haemoproteus species in Culicoides nubeculosus and to test if phylogenies based on the cytochrome b gene (cytb) reflect patterns of ookinete development in haemosporidian parasites. Additionally, one cytb lineage of Haemoproteus was identified to the species level and the in vitro gametogenesis and ookinete development of Haemoproteus hirundinis was characterised. Laboratory-reared C. nubeculosus were exposed by allowing them to take blood meals on naturally infected birds harbouring single infections of Haemoproteus belopolskyi (cytb lineage hHIICT1), Haemoproteus hirundinis (hDELURB2), Haemoproteus nucleocondensus (hGRW01) and Haemoproteus lanii (hRB1). Infected insects were dissected at intervals in order to detect sporogonic stages. In vitro exflagellation, gametogenesis and ookinete development of H. hirundinis were also investigated. Microscopic examination and PCR-based methods were used to confirm species identity. Bayesian phylogenetic inference was applied to study the relationships among Haemoproteus lineages. All studied parasites completed sporogony in C. nubeculosus. Ookinetes and sporozoites were found and described. Development of H. hirundinis ookinetes was similar both in vivo and in vitro. Developing ookinetes of this parasite possess long outgrowths, which extend longitudinally and produce the apical end of the ookinetes. A large group of closely related Haemoproteus species with a similar mode of ookinete development was determined. Bayesian analysis indicates that this character has phylogenetic value. The species identity of cytb lineage hDELURB2 was determined: it belongs to H. hirundinis. Culicoides nubeculosus is susceptible to and is a likely natural vector of numerous species of Haemoproteus parasites, thus worth attention in haemoproteosis epidemiology research. Data about in vitro development of haemoproteids provide valuable information about the rate of ookinete maturation and are recommended to use as helpful step during vector studies of haemosporidian parasites, particularly because they guide proper dissection interval of infected insects for ookinete detection during in vivo experiments. Additionally, in vitro studies readily identified patterns of morphological ookinete transformations, the characters of which are of phylogenetic value in haemosporidian parasites.
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Molecular characterization of six widespread avian haemoproteids, with description of three new Haemoproteus species.
Acta tropica, 2019Co-Authors: Gediminas Valkiūnas, Rasa Bernotienė, Dovilė Bukauskaitė, Mikas Ilgūnas, Carolina Romeiro Fernandes Chagas, Tanja Himmel, Josef Harl, Herbert Weissenböck, Tatjana A. IezhovaAbstract:Abstract Species of Haemoproteus (Haemosporida, Haemoproteidae) are widespread and often prevalent blood parasites of birds all over the word. They are particularly diverse in tropical countries. Due to limited knowledge of life cycles, these pathogens usually have been considered relatively benign and were neglected in veterinary medicine and bird management. However, recent molecular studies provided evidence that Haemoproteus parasites might cause severe diseases if they infect non–adapted (wrong) avian hosts due to marked damage of organs by exo-erythrocytic stages (megalomeronts). Additionally, high Haemoproteus infections are lethal to blood-sucking insects. Molecular markers are essential for reliable detection and species identification both at tissue stages in vertebrates and sporogonic stages in arthropods however, remain insufficiently developed for wildlife haemosporidian parasites. This study combined PCR-based and microscopic approaches and reported cytochrome b gene (cytb) and apicoplast gene (clpc) markers for characterization of six widespread species of haemoproteids parasitizing common birds wintering in tropics and subtropics of the Old World. Three new Haemoproteus species were described using morphological and molecular markers. Molecular characterization of haemoproteids parasitizing falcons was developed. Morphological and phylogenetic characterization of Haemoproteus tinnunculi (cytb lineage hFALSUB01), H. brachiatus (hLK03), H. parabelopolskyi (hSYAT1), H. homogeneae n. sp. (hSYAT16), H. homopicae n. sp. (hGAGLA07) and H. homominutus n. sp. (hCUKI1) was performed and provides clues for infections diagnostics. This study adds three species to the group of morphologically readily distinct Haemoproteus parasites, which differ in few base pairs (
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High susceptibility of the laboratory-reared biting midges Culicoides nubeculosus to Haemoproteus infections, with review on Culicoides species that transmit avian haemoproteids.
Parasitology, 2018Co-Authors: Dovilė Bukauskaitė, Tatjana A. Iezhova, Mikas Ilgūnas, Gediminas ValkiūnasAbstract:Haemosporidian parasites belonging to Haemoproteus cause avian diseases, however, vectors remain unidentified for the majority of described species. We used the laboratory-reared biting midges Culicoides nubeculosus to determine if the sporogonic development of three widespread Haemoproteus parasites completes in this insect. The midges were reared and fed on one common blackbird, white wagtail and thrush nightingale naturally infected with Haemoproteus minutus, Haemoproteus motacillae and Haemoproteus attenuatus, respectively. The engorged females were dissected in order to follow their sporogonic development. Microscopic examination was used to identify sporogonic stages. Bayesian phylogeny based on partial cytochrome b gene was constructed in order to determine phylogenetic relationships among Culicoides species-transmitted haemoproteids. All three parasites completed sporogony. Phylogenetic analysis placed Culicoides species transmitted haemoproteids in one well-supported clade, proving that such analysis readily indicates groups of dipteran insects transmitting avian haemoproteids. Available data show that 11 species of Culicoides have been proved to support complete sporogony of 18 species of avian haemoproteids. The majority of Culicoides species can act as vectors for many Haemoproteus parasites, indicating the low specificity of these parasites to biting midges, whose are globally distributed. This calls for control of haemoproteid infections during geographical translocation of infected birds.
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The widespread biting midge Culicoides impunctatus (Ceratopogonidae) is susceptible to infection with numerous Haemoproteus (Haemoproteidae) species.
Parasites & Vectors, 2017Co-Authors: Rita Žiegytė, Gediminas Valkiūnas, Rasa Bernotienė, Tatjana A. Iezhova, Mikhail Markovets, Andrey Mukhin, Vaidas PalinauskasAbstract:Haemoproteus parasites are widespread, and some species cause disease in wild and domestic birds. However, the insect vectors remain unknown for the majority of species and genetic lineages of avian Haemoproteus. This information is crucial for better understanding the biology of haemoproteids, the epidemiology of haemoproteosis, and the development of morphological characters of sporogonic stages in wildlife haemosporidian parasites. It remains unclear whether the specificity of Haemoproteus parasites for vectors is broad or the transmission of a given parasite can be restricted to a single or few species of vectors. The aim of this study was to examine the sporogonic development of four species of common European avian haemoproteids in the common biting midge Culicoides impunctatus. Wild-caught females of C. impunctatus were infected experimentally by allowing them to take blood meals on naturally infected Muscicapa striata, Cyanistes caeruleus, Ficedula hypoleuca and Motacilla flava harbouring mature gametocytes of Haemoproteus balmorali (genetic lineage hSFC9), H. majoris (hPARUS1), H. motacillae (hYWT1) and H. pallidus (hPFC1), respectively. Infected insects were collected, maintained under laboratory conditions and dissected daily in order to detect the development of ookinetes, oocysts and sporozoites. Microscopic examination and polymerase chain reaction based methods were used to detect the parasites. Bayesian analysis was applied to identify phylogenetic relationships among Haemoproteus lineages. All investigated parasites completed sporogony in C. impunctatus, indicating broad susceptibility of this biting midge for numerous Haemoproteus parasites. Ookinetes, oocysts and sporozoites were reported, described and compared morphologically. The investigated parasite species can be distinguished at the sporogony stage, particularly with regards to the morphology and rate of development of mature ookinetes. Analysis of data from the literature, and this study, shows that 12 genetically distantly related Haemoproteus parasites complete sporogony in C. impunctatus. Susceptibility of C. impunctatus is broad for Haemoproteus parasites, indicating that this biting midge is an important natural vector of numerous species of avian haemoproteids in Europe. Some Haemoproteus species can be readily distinguished using morphological characters of ookinetes and sporozoites, as well as the rate of ookinete development. These characters can be used for the identification of Haemoproteus species during sporogony in vectors, and are worth more attention in these parasite taxonomy studies at the species levels.
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Biting midges (Culicoides, Diptera) transmit Haemoproteus parasites of owls: evidence from sporogony and molecular phylogeny.
Parasites & vectors, 2015Co-Authors: Dovilė Bukauskaitė, Vaidas Palinauskas, Rasa Bernotienė, Tatjana A. Iezhova, Rita Žiegytė, Mikhail Markovets, Dimitar Dimitrov, Mikas Ilgūnas, Gediminas ValkiūnasAbstract:Background Haemoproteus parasites are widespread, and several species cause diseases both in birds and blood-sucking insects. These pathogens are transmitted by dipterans belonging to the Ceratopogonidae and Hippoboscidae, however certain vector species remain unknown for the majority of Haemoproteus spp. Owls are often infected by Haemoproteus parasites, but experimental studies on vectors of these infections are lacking. The aim of this study was to investigate sporogonic development of two widespread Haemoproteus parasites of owls, H. noctuae and H. syrnii in experimentally infected biting midges Culicoides impunctatus and Culicoides nubeculosus. We also followed in vitro sporogonic development of these infections and determined their phylogenetic relationships with Haemoproteus spp., for which vectors have been identified.
Ravinder N. M. Sehgal - One of the best experts on this subject based on the ideXlab platform.
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A fatal case of a captive snowy owl (Bubo scandiacus) with Haemoproteus infection in Japan.
Parasitology research, 2020Co-Authors: Megumu Yoshimoto, Kenichi Ozawa, Hirotaka Kondo, Yusuke Echigoya, Hisashi Shibuya, Yukita Sato, Ravinder N. M. SehgalAbstract:Parasites of the genus Haemoproteus are vector-borne avian haemosporidia commonly found in bird species of the world. Haemoproteus infections are typically considered relatively benign in birds. However, some Haemoproteus species cause severe disease and mortality, especially for captive birds removed from their original habitat. In September 2018, a captive 15-year-old snowy owl (Bubo scandiacus), kept in a zoological garden of Japan, died subacutely after presenting leg dysfunction. This case showed significantly low PCV and elevated AST, ALT, CK, and LDH values. Many megalomeronts with prominent morphological characteristics of Haemoproteus were observed in the left leg muscles. Those megalomeronts exhibited multilocular structures and were internally filled with merozoites. A new lineage of Haemoproteus was detected by subsequent PCR for the cytochrome b (cytb) gene of avian haemosporidia from DNA extracted from several organ tissues. The detected lineage was classified in the subgenus ParaHaemoproteus and was similar to those from the wild birds inhabiting the region including the study area, suggesting that this snowy owl likely acquired its infection from wild birds. This is the first report of a fatal case of a captive bird with a locally transmitted Haemoproteus infection in Japan. We considered the pathogenicity of this infection in conjunction with the clinical course and hematology results. We surmise that snowy owls may be particularly susceptible to infection with Haemoproteus parasites, and warming northern temperatures may exacerbate the overall health of these and other high latitude birds. Further research into the prevalence of Haemoproteus in wild birds near zoological gardens and potential biting midge vectors is necessary for the ex situ conservation of introduced birds.
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Delineation of the Genera Haemoproteus and Plasmodium Using RNA-Seq and Multi-gene Phylogenetics
Journal of molecular evolution, 2018Co-Authors: Jasper Toscani Field, Staffan Bensch, Gediminas Valkiūnas, Josh Weinberg, Nubia E. Matta, Ravinder N. M. SehgalAbstract:Members of the order Haemosporida are protist parasites that infect mammals, reptiles and birds. This group includes the causal agents of malaria, Plasmodium parasites, the genera Leucocytozoon and Fallisia, as well as the species rich genus Haemoproteus with its two subgenera Haemoproteus and ParaHaemoproteus. Some species of Haemoproteus cause severe disease in avian hosts, and these parasites display high levels of diversity worldwide. This diversity emphasizes the need for accurate evolutionary information. Most molecular studies of wildlife haemosporidians use a bar coding approach by sequencing a fragment of the mitochondrial cytochrome b gene. This method is efficient at differentiating parasite lineages but insufficient for accurate phylogenetic inferences in highly diverse taxa such as haemosporidians. Recent studies have utilized multiple mitochondrial genes (cyt b, cox1 and cox3), sometimes combined with a few apicoplast and nuclear genes. These studies have been highly successful with one notable exception: the evolutionary relationships of the genus Haemoproteus remain unresolved. Here we describe the transcriptome of Haemoproteus columbae and investigate its phylogenetic position recovered from a multi-gene dataset (600 genes). This genomic approach restricts the taxon sampling to 18 species of apicomplexan parasites. We employed Bayesian inference and maximum likelihood methods of phylogenetic analyses and found H. columbae and a representative from the subgenus ParaHaemoproteus to be sister taxa. This result strengthens the hypothesis of genus Haemoproteus being monophyletic; however, resolving this question will require sequences of orthologs from, in particular, representatives of Leucocytozoon species.
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Molecular characterization of Haemoproteus sacharovi (Haemosporida, Haemoproteidae), a common parasite of columbiform birds, with remarks on classification of haemoproteids of doves and pigeons.
Zootaxa, 2013Co-Authors: Asta Krizanauskiene, Staffan Bensch, Vaidas Palinauskas, Tatjana A. Iezhova, Jenny S. Carlson, Ravinder N. M. Sehgal, Gediminas ValkiunasAbstract:Haemoproteus (Haemosporida, Haemoproteidae) is the largest genus of avian haemosporidian parasites, some species of which cause lethal diseases in birds. Subgenera ParaHaemoproteus and Haemoproteus are usually accepted in this genus; these parasites are transmitted by biting midges (Ceratopogonidae) and hippoboscid flies (Hippoboscidae), respectively. As of yet, species of ParaHaemoproteus have not been reported to infect doves and pigeons (Columbiformes), parasites of these birds have not been reported to be transmitted by biting midges (Ceratopogonidae). Applying microscopy and PCR based methods, we identified mitochondrial cytochrome b (cyt b) sequences of Haemoproteus sacharovi, a widespread parasite of doves and pigeons. Phylogenetic relationships of dove haemoproteids, which traditionally have been classified in the subgenus Haemoproteus, showed that H. sacharovi and H. turtur, common parasites of doves, branch in the clade with ParaHaemoproteus species, indicating that these haemoproteids may belong to this subgenus and are likely transmitted by biting midges. This study provides barcodes for H. sacharovi, clarifies the taxonomic positions of H. sacharovi and H. turtur, and indicates directions for development of classification of avian haemoproteid species. Our analysis shows that the current subgeneric classification of avian haemoproteids is generally effective, but the position of some species may need to be revised.
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TWO NEW Haemoproteus SPECIES (HAEMOSPORIDA: HAEMOPROTEIDAE) FROM COLUMBIFORM BIRDS
The Journal of parasitology, 2012Co-Authors: Gediminas Valkiūnas, Tatjana A. Iezhova, Edward L. Evans, Jenny S. Carlson, Juan E. Martínez-gómez, Ravinder N. M. SehgalAbstract:Here we describe Haemoproteus (Haemoproteus) multivolutinus n. sp. from a tambourine dove (Turtur timpanistria )o f Uganda and Haemoproteus (Haemoproteus) paramultipigmentatus n. sp. (Haemosporida, Haemoproteidae) from the Socorro common ground dove (Columbina passerina socorroensis) of Socorro Island, Mexico. These parasites are described based on the morphology of their blood stages and segments of the mitochondrial cytochrome b gene that can be used for molecular identification and diagnosis of these species. Gametocytes of H. multivolutinus possess rod-like pigment granules and are evenly packed with volutin, which masks pigment granules and darkly stains both macro- and microgametocytes in the early stages of their development. Based on these 2 characters, H. multivolutinus can be readily distinguished from other species of hemoproteids parasitizing columbiform (Columbiformes) birds. Haemoproteus paramultipigmentatus resembles Haemoproteus multipigmentatus; it can be distinguished from the latter parasite primarily due to the broadly ovoid shape of its young gametocytes and significantly fewer pigment granules in its fully developed gametocytes. We provide illustrations of blood stages of the new species, and phylogenetic analyses identify DNA lineages closely related to these parasites. Cytochrome b lineages of Haemoproteus multivolutinus and H. paramultipigmentatus cluster with hippoboscid-transmitted lineages of hemoproteids; thus these parasites likely belong to the subgenus Haemoproteus .W e emphasize the importance of using cytochrome b sequences in conjunction with thorough microscopic descriptions to facilitate future identification of these and other avian hemosporidian species.
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New Avian Haemoproteus Species (Haemosporida: Haemoproteidae) From African Birds, with a Critique of the Use of Host Taxonomic Information in Hemoproteid Classification
The Journal of parasitology, 2011Co-Authors: Tatjana A. Iezhova, Ravinder N. M. Sehgal, Molly Dodge, Thomas B. Smith, Gediminas ValkiūnasAbstract:abstract: Haemoproteus (ParaHaemoproteus) micronuclearis n. sp., Haemoproteus (ParaHaemoproteus) nucleofascialis n. sp., Haemoproteus (ParaHaemoproteus) paranucleophilus n. sp., and Haemoproteus (ParaHaemoproteus) homobelopolskyi n. sp. (Haemosporida, Haemoproteidae) are described from African passeriform birds based on the morphology of their blood stages and segments of the mitochondrial cytochrome b gene. Red-billed quelea (Quelea quelea), red-headed malimbe (Malimbus rubricollis), and black-headed weaver (Ploceus melanocephalus) are the type vertebrate hosts of new hemoproteids. It is probable that new species have wide distribution in weavers in sub-Saharan Africa. Both H. micronuclearis and H. nucleofascialis can be readily distinguished from other avian hemoproteids by tiny, compact microgametocyte nuclei that are significantly smaller than macrogametocyte nuclei and are a rare character of hemosporidian parasites. Gametocytes of H. paranucleophilus are closely appressed to the erythrocyte nuclei ...
Rasa Bernotienė - One of the best experts on this subject based on the ideXlab platform.
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The use of nest boxes in determination of biting midges involved in transmission of Haemoproteus parasites
2020Co-Authors: Rita Žiegytė, Vaidas Palinauskas, Andrey Mukhin, Elena Platonova, Egidijus Kinderis, Rasa BernotienėAbstract:Abstract BackgroundCulicoides biting midges (Diptera, Ceratopogonidae) are known to be vectors of avian Haemoproteus parasites. These parasites cause disease and pathology in birds. The diversity of biting midges in Europe is great, but only four Culicoides species are known to be vectors of avian Haemoproteus parasites. In general, our knowledge about the role of the particular Culicoides species in transmission of Haemoproteus parasites remains insufficient. Information gaps hinder a better understanding of parasite biology and the epizootiology of parasite-caused diseases. The aim of this study was to determine new ornithophilic Culicoides species potentially involved in local transmission of Haemoproteus parasites. To do this we collected biting midges in bird nest boxes, identified their species and prevalence of haemoproteids in insects as well as in juvenile birds during breeding season.Methods Biting midges were collected from bird nest boxes, identified and parous females were tested individually for the presence of haemoproteids. The blood of juvenile birds was sampled to determine a local transmission of Haemoproteus spp. in the study area. We have used both microscopy and PCR-based methods.ResultsIn all, 293 Culicoides females belonging to 11 species were collected from nest boxes. Culicoides pictipennis, Culicoides segnis and Culicoides kibunensis were determined as dominant species collected using this method. Culicoides kibunensis was found to be infected with Haemoproteus lanii (genetic lineage hRB1), C.pictipennis and Culicoides punctatus− with Haemoproteus minutus (hTUPHI01 and hTURDUS2, respectively), C.segnis− with Haemoproteus majoris (hCWT4), H.minutus (hTURDUS2) and Haemoproteus tartakovskyi (hSISKIN1). From 187 studied juvenile birds 31 were infected with H. majoris (hCWT4, hPARUS1, hPHSIB1, hWW2) and Haemoproteus sp. (hPARUS10) parasites, which are widespread in Europe.Conclusions Our results provided information about the distribution of biting midge species and determined new ornithophilic Culicoides species at the study site. This study contributes to epizootiology of avian Haemoproteus infections by specifying Culicoides species that likely are responsible for the transmission of haemoproteids in Europe. Used method is suitable for better understanding vector ecology and evaluating the role of different blood sucking insects in transmission of haemoproteids in different wild ecosystems.
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Sporogony of four Haemoproteus species (Haemosporida: Haemoproteidae), with report of in vitro ookinetes of Haemoproteus hirundinis: phylogenetic inference indicates patterns of haemosporidian parasite ookinete development.
Parasites & vectors, 2019Co-Authors: Carolina Romeiro Fernandes Chagas, Rasa Bernotienė, Tatjana A. Iezhova, Dovilė Bukauskaitė, Mikas Ilgūnas, Gediminas ValkiūnasAbstract:Haemoproteus (ParaHaemoproteus) species (Haemoproteidae) are widespread blood parasites that can cause disease in birds, but information about their vector species, sporogonic development and transmission remain fragmentary. This study aimed to investigate the complete sporogonic development of four Haemoproteus species in Culicoides nubeculosus and to test if phylogenies based on the cytochrome b gene (cytb) reflect patterns of ookinete development in haemosporidian parasites. Additionally, one cytb lineage of Haemoproteus was identified to the species level and the in vitro gametogenesis and ookinete development of Haemoproteus hirundinis was characterised. Laboratory-reared C. nubeculosus were exposed by allowing them to take blood meals on naturally infected birds harbouring single infections of Haemoproteus belopolskyi (cytb lineage hHIICT1), Haemoproteus hirundinis (hDELURB2), Haemoproteus nucleocondensus (hGRW01) and Haemoproteus lanii (hRB1). Infected insects were dissected at intervals in order to detect sporogonic stages. In vitro exflagellation, gametogenesis and ookinete development of H. hirundinis were also investigated. Microscopic examination and PCR-based methods were used to confirm species identity. Bayesian phylogenetic inference was applied to study the relationships among Haemoproteus lineages. All studied parasites completed sporogony in C. nubeculosus. Ookinetes and sporozoites were found and described. Development of H. hirundinis ookinetes was similar both in vivo and in vitro. Developing ookinetes of this parasite possess long outgrowths, which extend longitudinally and produce the apical end of the ookinetes. A large group of closely related Haemoproteus species with a similar mode of ookinete development was determined. Bayesian analysis indicates that this character has phylogenetic value. The species identity of cytb lineage hDELURB2 was determined: it belongs to H. hirundinis. Culicoides nubeculosus is susceptible to and is a likely natural vector of numerous species of Haemoproteus parasites, thus worth attention in haemoproteosis epidemiology research. Data about in vitro development of haemoproteids provide valuable information about the rate of ookinete maturation and are recommended to use as helpful step during vector studies of haemosporidian parasites, particularly because they guide proper dissection interval of infected insects for ookinete detection during in vivo experiments. Additionally, in vitro studies readily identified patterns of morphological ookinete transformations, the characters of which are of phylogenetic value in haemosporidian parasites.
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Molecular characterization of six widespread avian haemoproteids, with description of three new Haemoproteus species.
Acta tropica, 2019Co-Authors: Gediminas Valkiūnas, Rasa Bernotienė, Dovilė Bukauskaitė, Mikas Ilgūnas, Carolina Romeiro Fernandes Chagas, Tanja Himmel, Josef Harl, Herbert Weissenböck, Tatjana A. IezhovaAbstract:Abstract Species of Haemoproteus (Haemosporida, Haemoproteidae) are widespread and often prevalent blood parasites of birds all over the word. They are particularly diverse in tropical countries. Due to limited knowledge of life cycles, these pathogens usually have been considered relatively benign and were neglected in veterinary medicine and bird management. However, recent molecular studies provided evidence that Haemoproteus parasites might cause severe diseases if they infect non–adapted (wrong) avian hosts due to marked damage of organs by exo-erythrocytic stages (megalomeronts). Additionally, high Haemoproteus infections are lethal to blood-sucking insects. Molecular markers are essential for reliable detection and species identification both at tissue stages in vertebrates and sporogonic stages in arthropods however, remain insufficiently developed for wildlife haemosporidian parasites. This study combined PCR-based and microscopic approaches and reported cytochrome b gene (cytb) and apicoplast gene (clpc) markers for characterization of six widespread species of haemoproteids parasitizing common birds wintering in tropics and subtropics of the Old World. Three new Haemoproteus species were described using morphological and molecular markers. Molecular characterization of haemoproteids parasitizing falcons was developed. Morphological and phylogenetic characterization of Haemoproteus tinnunculi (cytb lineage hFALSUB01), H. brachiatus (hLK03), H. parabelopolskyi (hSYAT1), H. homogeneae n. sp. (hSYAT16), H. homopicae n. sp. (hGAGLA07) and H. homominutus n. sp. (hCUKI1) was performed and provides clues for infections diagnostics. This study adds three species to the group of morphologically readily distinct Haemoproteus parasites, which differ in few base pairs (
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The widespread biting midge Culicoides impunctatus (Ceratopogonidae) is susceptible to infection with numerous Haemoproteus (Haemoproteidae) species.
Parasites & Vectors, 2017Co-Authors: Rita Žiegytė, Gediminas Valkiūnas, Rasa Bernotienė, Tatjana A. Iezhova, Mikhail Markovets, Andrey Mukhin, Vaidas PalinauskasAbstract:Haemoproteus parasites are widespread, and some species cause disease in wild and domestic birds. However, the insect vectors remain unknown for the majority of species and genetic lineages of avian Haemoproteus. This information is crucial for better understanding the biology of haemoproteids, the epidemiology of haemoproteosis, and the development of morphological characters of sporogonic stages in wildlife haemosporidian parasites. It remains unclear whether the specificity of Haemoproteus parasites for vectors is broad or the transmission of a given parasite can be restricted to a single or few species of vectors. The aim of this study was to examine the sporogonic development of four species of common European avian haemoproteids in the common biting midge Culicoides impunctatus. Wild-caught females of C. impunctatus were infected experimentally by allowing them to take blood meals on naturally infected Muscicapa striata, Cyanistes caeruleus, Ficedula hypoleuca and Motacilla flava harbouring mature gametocytes of Haemoproteus balmorali (genetic lineage hSFC9), H. majoris (hPARUS1), H. motacillae (hYWT1) and H. pallidus (hPFC1), respectively. Infected insects were collected, maintained under laboratory conditions and dissected daily in order to detect the development of ookinetes, oocysts and sporozoites. Microscopic examination and polymerase chain reaction based methods were used to detect the parasites. Bayesian analysis was applied to identify phylogenetic relationships among Haemoproteus lineages. All investigated parasites completed sporogony in C. impunctatus, indicating broad susceptibility of this biting midge for numerous Haemoproteus parasites. Ookinetes, oocysts and sporozoites were reported, described and compared morphologically. The investigated parasite species can be distinguished at the sporogony stage, particularly with regards to the morphology and rate of development of mature ookinetes. Analysis of data from the literature, and this study, shows that 12 genetically distantly related Haemoproteus parasites complete sporogony in C. impunctatus. Susceptibility of C. impunctatus is broad for Haemoproteus parasites, indicating that this biting midge is an important natural vector of numerous species of avian haemoproteids in Europe. Some Haemoproteus species can be readily distinguished using morphological characters of ookinetes and sporozoites, as well as the rate of ookinete development. These characters can be used for the identification of Haemoproteus species during sporogony in vectors, and are worth more attention in these parasite taxonomy studies at the species levels.
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Biting midges (Culicoides, Diptera) transmit Haemoproteus parasites of owls: evidence from sporogony and molecular phylogeny.
Parasites & vectors, 2015Co-Authors: Dovilė Bukauskaitė, Vaidas Palinauskas, Rasa Bernotienė, Tatjana A. Iezhova, Rita Žiegytė, Mikhail Markovets, Dimitar Dimitrov, Mikas Ilgūnas, Gediminas ValkiūnasAbstract:Background Haemoproteus parasites are widespread, and several species cause diseases both in birds and blood-sucking insects. These pathogens are transmitted by dipterans belonging to the Ceratopogonidae and Hippoboscidae, however certain vector species remain unknown for the majority of Haemoproteus spp. Owls are often infected by Haemoproteus parasites, but experimental studies on vectors of these infections are lacking. The aim of this study was to investigate sporogonic development of two widespread Haemoproteus parasites of owls, H. noctuae and H. syrnii in experimentally infected biting midges Culicoides impunctatus and Culicoides nubeculosus. We also followed in vitro sporogonic development of these infections and determined their phylogenetic relationships with Haemoproteus spp., for which vectors have been identified.
Vaidas Palinauskas - One of the best experts on this subject based on the ideXlab platform.
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The use of nest boxes in determination of biting midges involved in transmission of Haemoproteus parasites
2020Co-Authors: Rita Žiegytė, Vaidas Palinauskas, Andrey Mukhin, Elena Platonova, Egidijus Kinderis, Rasa BernotienėAbstract:Abstract BackgroundCulicoides biting midges (Diptera, Ceratopogonidae) are known to be vectors of avian Haemoproteus parasites. These parasites cause disease and pathology in birds. The diversity of biting midges in Europe is great, but only four Culicoides species are known to be vectors of avian Haemoproteus parasites. In general, our knowledge about the role of the particular Culicoides species in transmission of Haemoproteus parasites remains insufficient. Information gaps hinder a better understanding of parasite biology and the epizootiology of parasite-caused diseases. The aim of this study was to determine new ornithophilic Culicoides species potentially involved in local transmission of Haemoproteus parasites. To do this we collected biting midges in bird nest boxes, identified their species and prevalence of haemoproteids in insects as well as in juvenile birds during breeding season.Methods Biting midges were collected from bird nest boxes, identified and parous females were tested individually for the presence of haemoproteids. The blood of juvenile birds was sampled to determine a local transmission of Haemoproteus spp. in the study area. We have used both microscopy and PCR-based methods.ResultsIn all, 293 Culicoides females belonging to 11 species were collected from nest boxes. Culicoides pictipennis, Culicoides segnis and Culicoides kibunensis were determined as dominant species collected using this method. Culicoides kibunensis was found to be infected with Haemoproteus lanii (genetic lineage hRB1), C.pictipennis and Culicoides punctatus− with Haemoproteus minutus (hTUPHI01 and hTURDUS2, respectively), C.segnis− with Haemoproteus majoris (hCWT4), H.minutus (hTURDUS2) and Haemoproteus tartakovskyi (hSISKIN1). From 187 studied juvenile birds 31 were infected with H. majoris (hCWT4, hPARUS1, hPHSIB1, hWW2) and Haemoproteus sp. (hPARUS10) parasites, which are widespread in Europe.Conclusions Our results provided information about the distribution of biting midge species and determined new ornithophilic Culicoides species at the study site. This study contributes to epizootiology of avian Haemoproteus infections by specifying Culicoides species that likely are responsible for the transmission of haemoproteids in Europe. Used method is suitable for better understanding vector ecology and evaluating the role of different blood sucking insects in transmission of haemoproteids in different wild ecosystems.
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The widespread biting midge Culicoides impunctatus (Ceratopogonidae) is susceptible to infection with numerous Haemoproteus (Haemoproteidae) species.
Parasites & Vectors, 2017Co-Authors: Rita Žiegytė, Gediminas Valkiūnas, Rasa Bernotienė, Tatjana A. Iezhova, Mikhail Markovets, Andrey Mukhin, Vaidas PalinauskasAbstract:Haemoproteus parasites are widespread, and some species cause disease in wild and domestic birds. However, the insect vectors remain unknown for the majority of species and genetic lineages of avian Haemoproteus. This information is crucial for better understanding the biology of haemoproteids, the epidemiology of haemoproteosis, and the development of morphological characters of sporogonic stages in wildlife haemosporidian parasites. It remains unclear whether the specificity of Haemoproteus parasites for vectors is broad or the transmission of a given parasite can be restricted to a single or few species of vectors. The aim of this study was to examine the sporogonic development of four species of common European avian haemoproteids in the common biting midge Culicoides impunctatus. Wild-caught females of C. impunctatus were infected experimentally by allowing them to take blood meals on naturally infected Muscicapa striata, Cyanistes caeruleus, Ficedula hypoleuca and Motacilla flava harbouring mature gametocytes of Haemoproteus balmorali (genetic lineage hSFC9), H. majoris (hPARUS1), H. motacillae (hYWT1) and H. pallidus (hPFC1), respectively. Infected insects were collected, maintained under laboratory conditions and dissected daily in order to detect the development of ookinetes, oocysts and sporozoites. Microscopic examination and polymerase chain reaction based methods were used to detect the parasites. Bayesian analysis was applied to identify phylogenetic relationships among Haemoproteus lineages. All investigated parasites completed sporogony in C. impunctatus, indicating broad susceptibility of this biting midge for numerous Haemoproteus parasites. Ookinetes, oocysts and sporozoites were reported, described and compared morphologically. The investigated parasite species can be distinguished at the sporogony stage, particularly with regards to the morphology and rate of development of mature ookinetes. Analysis of data from the literature, and this study, shows that 12 genetically distantly related Haemoproteus parasites complete sporogony in C. impunctatus. Susceptibility of C. impunctatus is broad for Haemoproteus parasites, indicating that this biting midge is an important natural vector of numerous species of avian haemoproteids in Europe. Some Haemoproteus species can be readily distinguished using morphological characters of ookinetes and sporozoites, as well as the rate of ookinete development. These characters can be used for the identification of Haemoproteus species during sporogony in vectors, and are worth more attention in these parasite taxonomy studies at the species levels.
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Biting midges (Culicoides, Diptera) transmit Haemoproteus parasites of owls: evidence from sporogony and molecular phylogeny.
Parasites & vectors, 2015Co-Authors: Dovilė Bukauskaitė, Vaidas Palinauskas, Rasa Bernotienė, Tatjana A. Iezhova, Rita Žiegytė, Mikhail Markovets, Dimitar Dimitrov, Mikas Ilgūnas, Gediminas ValkiūnasAbstract:Background Haemoproteus parasites are widespread, and several species cause diseases both in birds and blood-sucking insects. These pathogens are transmitted by dipterans belonging to the Ceratopogonidae and Hippoboscidae, however certain vector species remain unknown for the majority of Haemoproteus spp. Owls are often infected by Haemoproteus parasites, but experimental studies on vectors of these infections are lacking. The aim of this study was to investigate sporogonic development of two widespread Haemoproteus parasites of owls, H. noctuae and H. syrnii in experimentally infected biting midges Culicoides impunctatus and Culicoides nubeculosus. We also followed in vitro sporogonic development of these infections and determined their phylogenetic relationships with Haemoproteus spp., for which vectors have been identified.
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Haemoproteus infections (Haemosporida, Haemoproteidae) kill bird-biting mosquitoes
Parasitology research, 2013Co-Authors: Gediminas Valkiūnas, Vaidas Palinauskas, Rasa Bernotienė, Dovilė Bukauskaitė, Rita Kazlauskienė, Tatjana A. IezhovaAbstract:Haemoproteus parasites (Haemosporida, Haemoproteidae) are widespread; some species cause severe diseases in avian hosts. Heavy Haemoproteus infections are often lethal for biting midges (Ceratopogonidae), which transmit avian haemoproteids, but there is no information regarding detrimental effect on other blood-sucking insects. We examined effects of Haemoproteus tartakovskyi (lineage hSISKIN1), Haemoproteus lanii (lineages hRB1and hRBS2) and Haemoproteus balmorali (lineage hCOLL3) on the survival of Ochlerotatus cantans, a widespread Eurasian mosquito. Wild-caught females were infected by allowing them to feed on naturally infected birds with light (0.01 %) and high (3.0–9.6 %) parasitaemia. Mosquitoes fed on uninfected birds were used as controls. Both experimental and control groups were maintained under the same laboratory conditions until 20 days post-exposure (dpe). Dead insects were counted daily and used for parasitological examination and PCR-based testing. No difference was discernible in the survival rate of control mosquitoes and those fed on meal with light parasitaemia. There was a highly significant difference in the survival rate between the control group and all groups fed on meals with high parasitaemia, with the greatest mortality reported 1–3 dpe. For 4 dpe, the percentage of survived control mosquitoes (88 %) was 2.2-, 3.6- and 4-fold greater than that of groups fed on meals with high parasitaemia of H. balmorali, H. tartakovskyi and H. lanii, respectively. Numerous ookinetes were observed in the gut area and adjacent tissues located in the head, thorax and abdomen of infected insects 0.5–1 dpe. The migrating parasites damage organs throughout the entire body of mosquitoes; that is the main reason of mortality. To the end of this study, 46 % of mosquitoes survived in control group, but the survival rates of experimental mosquitoes fed on meals with high parasitaemia were between 2.6- and 5.8-fold lower. This study indicates that widespread Haemoproteus infections are markedly virulent for bird-biting mosquitoes, which rapidly die after feeding on heavily infected blood meals.
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Molecular characterization and distribution of Haemoproteus minutus (Haemosporida, Haemoproteidae): A pathogenic avian parasite
Parasitology international, 2013Co-Authors: Vaidas Palinauskas, Staffan Bensch, Tatjana A. Iezhova, Mikhail Markovets, Asta Križanauskienė, Gediminas ValkiūnasAbstract:Recently, the lineage hTURDUS2 of Haemoproteus minutus (Haemosporida, Haemoproteidae) was reported to cause mortality in captive parrots. This parasite lineage is widespread and prevalent in the blackbird Turdus merula throughout its entire distribution range. Species identity of other closely related lineages recently reported in dead parrots remains unclear, but will be important to determine for a better understanding of the epidemiology of haemoproteosis. Using polymerase chain reaction (PCR)-based and microscopic methods, we analyzed 265 blood samples collected from 52 species of wild birds in Eurasia (23 samples from Kamchatka Peninsula, 73 from Sakhalin Island, 150 from Ekaterinburg and 19 from Irkutsk regions of Russia). Single infections of the lineages hTURDUS2 (hosts are redwing Turdus iliacus and fieldfare Turdus pilaris), hTUPHI1 (song thrush Turdus philomelos) and hTUCHR01 (fieldfare, redwing, song thrush and brown-headed thrush Turdus chysolaus) were detected. We identified species of these haemoproteids based on morphology of their blood stages and conclude that these lineages belong to H. minutus, a widespread parasite of different species of thrushes (genus Turdus), which serve as reservoir hosts of this haemoproteid infection. Phylogenetic analysis shows that the lineages hTURDUS2, hTUCHR01 and hTUPHI1 of H. minutus are closely related to Haemoproteus pallidus (lineages hPFC1 and hCOLL2), Haemoproteus pallidulus (hSYAT03), and Haemoproteus sp. (hMEUND3); genetic distance among their mitochondrial cytochrome b (cyt b) lineages is small (
