Keys to the avian malaria parasites.

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malaria 40 Title: Malaria JournalKeys to the avian malaria parasitesGediminas ValkiūnasTatjana A. IezhovaPublication date (epub): 5/2018Publication date (pmc-release):
malaria 373 birds. These pathogens cause pathology of blood and various organs, often resulting in severe avian malaria . Numerous recent studies have reported DNA sequences of avian malaria parasites, indicating rich genetic
malaria 443 often resulting in severe avian malaria. Numerous recent studies have reported DNA sequences of avian malaria parasites, indicating rich genetic diversity and the possible existence of many undescribed species.
malaria 916 described. However, keys for their identification are unavailable or incomplete. Identification of avian malaria parasites remains a difficult task even for experts, and this precludes development of avian malariology,
malaria 1077 and this precludes development of avian malariology, particularly in wildlife. Here, keys for avian malaria parasites have been developed as a baseline for assisting academic and veterinary medicine researchers
malaria 2572 markers are unavailable for 58% of described Plasmodium parasites, raising a task for the current avian malaria researchers to fill up this gap.BackgroundMalaria parasites of the genus Plasmodium (Haemosporida, Plasmodiidae)
malaria 2752 Plasmodium (Haemosporida, Plasmodiidae) inhabit all major groups of terrestrial vertebrates. Avian malaria parasites is a peculiar group among them, particularly due to the ability of numerous species to develop
malaria 3315 Psorophora) for completing sporogony and transmission [[1], [8]–[11]]. This is not the case in mammalian malaria parasites whose are transmitted mostly by Anopheles species [[1], [12]–[14]]. Furthermore, sporogony
malaria 3643 [[1], [8], [15], [16]]. These features likely contributed to the global distribution of some avian malaria infections, which are actively transmitted in countries with warm and cold climates, including regions
malaria 3821 cold climates, including regions close to the Polar Circles [[6], [17]–[19]].Life cycles of avian malaria parasites are similar in their basic features to those of human and other mammal Plasmodium species
malaria 4752 also have a broad range of avian hosts [[6], [8], [21]–[23]]. Erythrocytic merozoites of many avian malaria parasites can induce secondary tissue merogony in birds [[24], [25]]. The exo-erythrocytic merogony
malaria 5239 oocysts to the mosquito midgut wall [[27]]. These and some other features are not characteristics of malaria parasites of mammals, and this is reflected in genetic differences between these groups of parasites
malaria 5762 [[1], [13], [14], [34]]. While available evidence still supports this view for the primate and rodent malaria l parasites, there is increasing evidence that the pathogenicity of tissue stages of avian species of
malaria 5945 stages of avian species of Plasmodium has been significantly underestimated [[25]]. Even more, avian malaria is often a more severe disease than human malaria. There is recent experimental evidence of unexpected
malaria 5995 significantly underestimated [[25]]. Even more, avian malaria is often a more severe disease than human malaria . There is recent experimental evidence of unexpected pathology associated with obstructive development
malaria 6770 words, vertebrate host identity cannot be used as a taxonomic feature during identification of avian malaria parasites [[1], [12], [42]]. This raises questions about parasite species identification if the same
malaria 6975 same pathogen is found in unusual avian hosts. Molecular characterization is helpful in diagnosis of malaria infections, and has been developed for detection of some avian Plasmodium species [[21], [40]]. Molecular
malaria 7710 not been developed, and currently are difficult to develop due to significant genetic diversity of malaria parasites, which remain undescribed in wildlife. Morphological identification using microscopic examination
malaria 7862 wildlife. Morphological identification using microscopic examination of blood films remains important in malaria diagnostics in the wild, and is particularly valuable if it is applied in parallel with polymerase chain
malaria 8534 information.The main aim of this review is to develop easy-to-use keys for identification of avian malaria parasites using morphological features of their blood stages as a baseline for assisting academic and
malaria 9200 Plasmodium species detection and comparison was also summarized. This review might be helpful for wildlife malaria and veterinary medicine researchers aiming identification of avian malaria infections.MethodsFull-length
malaria 9275 be helpful for wildlife malaria and veterinary medicine researchers aiming identification of avian malaria infections.MethodsFull-length papers with descriptions of new Plasmodium species published in peer-reviewed
malaria 13064 Plasmodiidae (a–c), Garniidae (d–f), Haemoproteidae (g) and Leucocytozoidae (h, i). Note presence of malaria l pigment in species of Plasmodiidae (a–c) and Haemoproteidae (g) and its absence in species of Garniidae
malaria 17035 inhabit red blood cells and do not possess pigment granules in their blood stages (Fig. 1d–f).When malaria parasites of the Plasmodium genus are reported in blood films, the next step is to distinguish subgenera
malaria 19077 (Fig. 3p–r)…………………………………………… NovyellaeMain taxonomic characters of subgenera of avian malaria parasites [[8]]aExo-erythrocytic merogony takes place in cells of the haemopoietic system. Erythrocytic
malaria 21522 grown gametocytes are elongated (Fig. 4c–r). Pedunculated oocysts are absentWhen the subgenus of a malaria parasite has been identified, the next step is the species identification using the keys to species
malaria 50349 three main groups of obstacles, which a researcher usually faces during morphological identification of malaria parasites using microscopic examination of blood samples collected in the field. First, the quality
malaria 51040 Second, Plasmodium species parasitaemia is often light in natural infections in the wild. In other words, malaria parasites might be reported in blood films, but not all stages, which are needed for parasite species
malaria 51726 [56]]. These obstacles strengthen the need for the development of molecular characterization in avian malaria diagnostics, which is still only available for 44% of described parasite species, whose validity is
malaria 55373 of this subgenus have been characterized molecularly). Lack of molecular markers for many described malaria pathogens [[51], [53], [54], [56], [57], [59], [65]] precludes biodiversity research on Plasmodium species
malaria 55511 [56], [57], [59], [65]] precludes biodiversity research on Plasmodium species and recognition of new malaria pathogens, for whose detection, detailed comparison with already described and genetically characterized
malaria 55827 important task of current avian malariology (Table 7).This study shows that 55 described species of avian malaria parasites can be readily distinguished (Tables 3, 4, 5, 6, 7). Among them, 12, 16, 22, 4 and 1 species
malaria 56323 indicating that transmission of these pathogens occurs mainly in countries with warm climates. Those malaria parasites, which have adapted for transmission globally and have become cosmopolitan, are exceptions.
malaria 56919 Novyella are particularly diverse (Table 5). They represent approximately 40% of all described avian malaria pathogens, and 78% of Plasmodium species, which were discovered during past 15 years. Novyella parasites
malaria 57442 Novyella infections globally are unclear. Novyella species are the most poorly studied group of avian malaria pathogens, with nearly no information available about exo-erythrocytic development, virulence, sporogony
malaria 58032 information indicates that some Novyella species (P. ashfordi, P. rouxi) may cause severe and even lethal malaria in some birds due to blood pathology [[1], [8], [74], [75]], but the complete mechanism of their pathogenicity
malaria 58370 cycles and virulence of infections caused by Novyella species is an important task in current avian malaria research.Many species of Plasmodium inhabit numerous species of birds and use mosquitoes of different
malaria 58739 and strain varieties. Because of these morphological variants, it has been conventional in old avian malaria research (approximately between 1927 and 1995) that any new Plasmodium species description should only
malaria 60120 and were not accompanied with molecular characterization. Due to the huge genetic diversity of avian malaria pathogens and numerous genetic lineages reported in birds, some of these names might be validated in
malaria 65134 the original descriptions truly belong to corresponding species.Table 9List of species names of bird malaria parasites belonging to the categories of nomen nudum, nomen dubium, species inquirenda and incertae
malaria 66092 lagopi [[156]]Species inquirendaP. lairdi [[157]]Nomen nudumP. lenoblei [[56]]bSpecies inquirendaP. malaria e raupachi [[158]]Incertae sedisP. manwelli [[159]]Nomen nudumP. ninoxi [[160]]dSpecies inquirendaP.
malaria 68585 to different subgenera are common in wildlife, and the described cases of co-infections with several malaria parasites are not unpredictable [[45]]. However, description of new species from such co-infections
malaria 69415 avian hosts, calling for careful application of minor differences in blood stage morphology in avian malaria parasite taxonomy, particularly during co-infectionscBased on available information [[53]], P. buteonis
malaria 75043 were designated as holotypes. Single cells usually do not reflect entire morphological diversity of malaria parasites, so deposition of parahapantotype material is preferable in wildlife haemosporidian research
malaria 78858 them to avoid taxonomic confusion [[78]].The subgenus Papernaia was created for Novyella-like avian malaria parasites, whose erythrocytic meronts do not possess globules (Fig. 3f, h–l), structures of unclear
malaria 79101 [80]]. The feature of the presence or absence of such globules is used in distinguishing some species of malaria parasites belonging to subgenus Novyella during natural infections (Table 5). It is interesting to
malaria 80263 subsequent passages in siskins (Fig. 6c, d) illustrate this change. These experimental data indicate that malaria parasites which do not possess globules in natural hosts might develop this structure after artificial
malaria 82450 categories species inquirenda and incertae sedis should be considered in future taxonomic work of avian malaria parasite at species level. The majority of described Plasmodium parasites have not been characterized
malaria 82642 been characterized using molecular markers, which development is an essential task for current avian malaria researchers

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