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Table 9 List of species names of bird malaria parasites belonging to the categories of nomen nudum, nomen dubium, species inquirenda and incertae sedis

From: Keys to the avian malaria parasites

Name and references

Statusa

Plasmodium alaudae [57, 126]b

Species inquirenda

P. arachnidi [147]

Species inquirenda

P. bambusicolai [148]

Species inquirenda

P. beaucournui [56]b

Species inquirenda

P. bigueti [65]b

Species inquirenda

P. buteonis [53]c

Species inquirenda

P. coggeshalli [149]b

Species inquirenda

P. conturnixae [150]

Nomen nudum

P. corradettii [151]

Nomen dubium

P. danilewskyi [152, 153]

Incertae sedis

Plasmodium dherteae [56]b

Species inquirenda

P. gallinulae [130]

Incertae sedis

P. gambeli [3]

Nomen nudum

P. ghadiriani [56]b

Species inquirenda

P. golvani [56]b

Species inquirenda

P. herodiadis [130]

Species inquirenda

P. holti [154]

Nomen nudum

P. jiangi [155]

Species inquirenda

P. jeanriouxi [57]b

Species inquirenda

P. lagopi [156]

Species inquirenda

P. lairdi [157]

Nomen nudum

P. lenoblei [56]b

Species inquirenda

P. malariae raupachi [158]

Incertae sedis

P. manwelli [159]

Nomen nudum

P. ninoxi [160]d

Species inquirenda

P. noctuae [3, 126]

Species inquirenda

P. pachysomum [54]e

Species inquirenda

P. papernai [149]b

Species inquirenda

Plasmodium pfefferi [54]e

Species inquirenda

P. praecox [152]

Nomen nudum

P. reniai [57]b

Species inquirenda

P. rousseloti [161]

Species inquirenda

P. rouxi, as published in [54]f

Species inquirenda (probably a new Plasmodium species)

P. sergentorum [54]e

Species inquirenda

P. snounoui [56]b

Species inquirenda

P. spartani [162]

Nomen nudum

P. stellatum [54]e

Species inquirenda

P. struthionis [163]

Incertae sedis

P. tranieri [56]b

Species inquirenda

P. venkataramiahii [164]

Nomen nudum

  1. aNomenclature status of the species names published before 2000 was justified in [8]
  2. bPlasmodium beaucournui, P. bigueti, P. coggeshalli, P. dherteae, P. ghadiriani, P. golvani, P. jeanriouxi, P. lenoblei, P. papernai, P. reniai, P. snounoui, P. tranieri were named and described, and P. alaudae was re-described from individual birds co-infected with parasites belonging to subgenera Haemamoeba, Giovannolaia and Novyella [56, 57, 65, 149]. The authors of the original descriptions have grouped the blood stages visible in blood films and attributed them to different species provisionally, which is particularly obvious in case of parasites with elongate gametocytes. This makes species description and validation of parasite names questionable. Only single cells (erythrocytic meronts) were selected as holotypes in these parasite descriptions. However, due to morphological variation of blood stages of Plasmodium and presence of parasites at different stages of growth in each blood film, such methodology of designation of the type material can work only in case of exceptionally distinctive cell characters, which is not the case in all these parasite descriptions, particularly belonging to subgenus Haemamoeba. Molecular characterization of all these parasites is unavailable. It is clear from the original descriptions, that many individual birds were infected by representatives of several subgenera. However, the reported blood stages were selected and attributed to certain species without providing convincing explanations, making identifications difficult or even impossible based on available information. Co-infections of Plasmodium parasites belonging 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 hardly possible if the unique morphological characters of blood stages are absent, which is the case with P. beaucournui, P. bigueti, P. coggeshalli, P. dherteae, P. ghadiriani, P. golvani, P. jeanriouxi, P. lenoblei, P. papernai, P. reniai, P. snounoui, P. tranieri and also in re-description of P. alaudae. These parasites are considered as species inquirenda. Recent molecular studies provided molecular markers for distinguishing blood stages of Plasmodium species (Table 7). Examination of blood films from experimental infections shows variations in morphological characters of same parasite lineages in different avian hosts, calling for careful application of minor differences in blood stage morphology in avian malaria parasite taxonomy, particularly during co-infections
  3. cBased on available information [53], P. buteonis cannot be distinguished from P. circumflexum and other similar parasites of Giovannolaia (Plasmodium gabaldoni, Plasmodium homocircumflexum). The main feature, which has been noted to distinguish P. buteonis from P. circumflexum in the original description [53], is the presence up to 36 nuclei in mature erythrocytic meronts of the former. Plasmodium circumflexum produce less number of nuclei in mature meronts. However, the description of P. buteonis is based on high parasitemia (6.6%), with numerous multiple infections of the same erythrocytes, so it is difficult to rule out that 2 mature meronts were present in same cell in case of so great number of merozoites. Additionally, parasite morphology often changes during high parasitemia, so such samples should be carefully used in taxonomical descriptions. Plasmodium buteonis might be a valid name, but more research is needed to prove its validity. Molecular characterization of this parasite is absent, but is essential to solve the question about its validity
  4. dPlasmodium ninoxi was described from owl Ninox scutulata in co-infection with Haemoproteus sp. [160]. Only one erythrocyte with 2 binuclear growing meronts was detected; no other data about merogony in the blood were provided. Plasmodium ninoxi gametocytes were reported to be rounded. Based on available information, it seems that infected blood was exposed to air, which stimulated rounding-up of haemoproteid gametocytes [8], which were attributed to P. ninoxi. DNA sequence was provided (AY099035.1), and it belongs to Plasmodium sp. Plasmodium ninoxi description is incomplete. Re-description is needed, and it is possible due to available sequence information. The most similar cytb sequence belong to P. gallinaceum, P. relictum and P. circumflexum
  5. eDescriptions of P. pachysomum, P. pfefferi, P. sergentorum, P. stellatum [54] are incomplete. Information about morphology of gametocytes is absent. Molecular characterization is unavailable. Species identification is questionable based on the available information
  6. fPaperna et al. [54] published re-description of P. rouxi from non-type avian host (Alauda arvensis, Alaudidae instead of Passer hispaniolensis, Passeridae whose is the type host). The re-description is based on samples, which were collected beyond of the type locality (France, instead of Algeria which is the type locality). This contradicts the Article 75.3.6 of the International Code of Zoological Nomenclature [78]. Additionally, according to [54], the erythrocytic meronts of the parasite from A. arvensis do not possess refractive globules and gametocytes possess few tiny pigment granules (Figs. 8, 9 in [54]). These are not characters of P. rouxi, which was described by Sergent et al. [111]. Sergent’s original material from Algiers labelled “2198, 26.4.28, Institut Pasteur d’Algérie” is available in the Natural History Museum, London. Examination of this blood film showed that numerous erythrocytic meronts of this parasite possess refractive globules (Fig. 3f, j) and gametocytes possess few large (Fig. 4o) pigment granules. The latter character is an important feature of P. rouxi. Based on available information, the parasite described in [54] as ‘P. rouxi’ cannot be attributed to P. rouxi and is considered as a species inquirenda. The parasite described by Paperna et al. [54] is characterized by presence of (1) the relatively prominent cytoplasm in growing meronts and (2) tiny size of pigment granules in gametocytes, so might belong to a new Plasmodium species. Additional investigation is needed to answer this question. In the same study, Paperna et al. [54] described a new species Plasmodium mohammedi, which was reported, Passer domesticus (the common host of P. rouxi in Mediterranian region [68]). Blood stages of P. mohammedi are indistinguishable from P. rouxi [111], particularly due to the presence of refractive globules in erythrocytic meronts and large pigment granules in gametocytes (see Figs. 18–21 in [54]). Plasmodium mohammedi is a synonym of P. rouxi. Molecular identification of P. rouxi (lineage pPADOM16) was developed [68]. Application of the barcoding indicates that the details of disposition of nuclei in erythrocytic meronts during different infections, particularly in different avian hosts, is variable in P. rouxi, but binuclear “bow-tie” form parasites often are present (Fig. 3f) and can be used for this parasite species identification. Additionally, presence of few large pigment granules in mature gametocytes also is a characteristic feature, and it recommended to use for distinguishing P. rouxi infection (Fig. 4o) from other Novyella parasites producing tetranuclear erythrocytic meronts
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Malaria Journal

ISSN: 1475-2875

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