Overview

Brief Summary

There are more than 150 named species of Plasmodium that infect various species of vertebrates. Four species are well known as true parasites of humans, utilizing humans almost exclusively as a natural intermediate host in which they cause malaria: P. falciparum, P. vivax, P. ovale and P. malariae. In recent years it has become apparent that the simian malaria parasite P. knowlesi also regularly infects humans, as well as its natural monkey intermediate hosts. (Centers for Disease Control Parasites and Health website) Furthermore, it is now apparent that there are likely two distinct Plasmodium species that have both been referred to as P. ovale (Sutherland et al. 2010; Oguike et al. 2011).

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© Shapiro, Leo

Source: EOL Rapid Response Team

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Introduction

Organisms that belong to the genus Plasmodium are obligate eukaryotic parasites, best known as the etiological agent of human malaria. There are four parasites that infect humans and cause malaria: P. falciparum, P. vivax, P. malariae, and P. ovale. Although Plasmodium parasites infect a variety of vertebrate hosts (including primates, rodents, ungulates, birds, and lizards), they rarely cause severe disease in any vertebrate hosts other than humans. The most virulent of human parasites, P. falciparum and P. vivax (White, 2003), cause 300-500 million cases of debilitating or fatal disease worldwide.

Our understanding of the evolution and systematics of malaria parasites has changed significantly over the past 20 years with recent work (Perkins and Schall, 2002; Martinsen et al., 2008) indicating that the genus Plasmodium may not to be monophyletic, and includes parasites of other genera including Hepatocystis.

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Comprehensive Description

<em>Plasmodium</em> Life-cycle

Plasmodium parasites have an elaborate life-cyle with multiple stages:

  1. Infective stage, when the parasite enters the vertebrate host with a vector bite. This life stage is known as sporozoite.
  2. Exoerythrocytic stage, in which the sporozoite undergoes multiple rounds of asexual divisions (merogony or schizongony) and matures into merozoites.
  3. Erythrocytic stages, during which the organisms enter red blood cells (as merozoites), transform into the feeding stages (trophozoites), and then divide asexually into multiple new merozoites (schizont stage). During the schizont stage, some parasites differentiate into the reproductive forms (gametocytes) rather than the invasive merozoites. Gametocytes are classified as microgametocytes (that will become male gametes) and macrogametocytes (that will become female gametes). The gametocytes must mature through five stages before they become infective to the mosquito.

    Left: P. vivax in early trophozoite "ring" stage. Center: P. vivax, schizont stage. Right: P. falciparum, mature macrogametocyte.

  4. Reproductive stages, these begin when the vector takes a blood meal from the vertebrate host that contains mature gametocytes. In the vector the gametocytes transform into male and female gametes and merge to become a zygote (the only diploid stage in the organism's life-cycle). The zygote becomes an ookinate which invades the tissues of the vector midgut to become an oocyst. When the oocyst ruptures thousands of sporozoites emerge and travel to the vector's salivary glands, as it is through the saliva that they will enter the next vertebrate host.

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Hosts and Geographic Distribution of Some Parasites

ParasiteHostGeographic Distribution
Plasmodium reichenowiChimapanzeeAfrica
Plasmodium falciparumHuman Africa, Asia, South/Central America
Plasmodium fieldiMacaqueSoutheast Asia
Plasmodium simiovaleMacaqueSoutheast Asia
Plasmodium hylobatiMacaqueSoutheast Asia
Plasmodium inuiMacaqueSoutheast Asia
Plasmodium knowlesiMacaqueSoutheast Asia
Plasmodium coatneyiMacaqueSoutheast Asia
Plasmodium simium Spider MonkeySouth America
Plasmodium vivaxHumanAfrica, Asia, South/Central America
Plasmodium cynomolgiMacaqueSoutheast Asia
Plasmodium gonderiMadrillAfrica
Plasmodium malariaeHumanAfrica, Asia, South/Central America
Plasmodium brasilianumSpider/Howler/Night MonkeySouth America
Plasmodium ovaleHumanAfrica
Hepatocystis sp.Bat/PrimateAfrica, Asia
Plasmodium atheruriRodentAfrica
Plasmodium vinkeiRodentAfrica
Plasmodium chabaudiRodentAfrica
Plasmodium bergheiRodentAfrica
Plasmodium yoeliiRodentAfrica
Plasmodium mexicanumLizardNorth America
Plasmodium chiricahuaeLizardNorth America
Plasmodium elongatumBirdWorldwide
Plasmodium gallinaceumBirdSoutheast Asia
Plasmodium relictumBirdWorldwide
Plasmodium floridense
Lizard
Carribbean/Central America
Plasmodium azurophilum
Lizard
Carribbean/Central America
Plasmodium faichildi
Lizard
Central America
Plasmodium agamae
Lizard Africa
Plasmodium gigantum
Lizard Africa
Plasmodium mackerassaeLizardAustralia

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Characteristics

Organisms of the genus Plasmodium are defined as distinct from other Apicomplexa, and other organisms sometimes considered malaria parasites (Peréz-Tris et al. 2005), as parasites that both undergo merogony (multiple divisions of the nucleus followed by segmentation of the cytoplasm producing daughter cells called merozoites) in erythrocytes, and that produce hemozoin pigment, the crystalline by-product of hemoglobin digestion. Other members of the order Haemosporida vary in the combination of these characters. The family Haemoproteidae produces pigment, but merogony occurs in tissues other than erythrocytes. The families Garniidae and Leucocytozoidae do not produce pigment at any stages, but in the case of the former do show merogony in blood cells.

P. falciparum schizont stage parasite (center) showing merogony, recognizable by approximately 24 new merozoites identified by the Giemsa-stained nuclei (purple), and showing hemozoin pigment (yellow)

All malaria parasites have a sexual life stage that occurs in a blood-feeding insect, which is the definitive host for these organisms (also known as the “vector,” in epidemiology).

Traditionally, Plasmodium species were described based on morphological and morphometric characteristics, primarily of the blood stage of the lifecycle. Other characters used in the past for classification have been virulence-level and life-cycle period in the erythrocytic stage (24, 48, or 72 hours). Morphological characterization has often proven to be insufficient to distinguish species, and has also been shown to conflict with molecular sequence data. Scientists are beginning to use advanced methods and molecular characters for parasite taxonomy and phylogenetics. (See Discussion of Phylogenetic Relationships.)

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Description of Plasmodium

Plasmodiid apicomplexa, causative agent of malaria, gamonts in erythrocytes; merogony in erythrocytes and other tissues depending on species; hemozoin deposited; vectors mosquitoes or phlebotomine flies; in mammals, birds, and reptiles; cause of estimated two million deaths annually; most important human species are P. vivax (causes benign tertian malaria) and P. falciparum (causesmalignant tertian malaria); Type species P. malariae
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biopedia

Source: BioPedia

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Evolution and Systematics

Evolution

Discussion of Phylogenetic Relationships

View Plasmodium Tree

Phylogeny based on Escalante et al., 1995; Perkins and Schall, 2002; Vargas-Serrato et al., 2003; Martinsen et al., 2008.

The field of Plasmodium phylogenetics has been particularly dynamic over the past decade, and continues to be so as more species are added to the tree and older hypotheses are called into question.

The study of the molecular phylogenetics of malaria parasites began in 1991 (Waters et al., 1991), and the field continues to grow. The conclusions of many earlier studies were overturned later, as they were affected by difficulties stemming from insufficient taxon sampling and problems with the genes chosen for analysis. The choice of both the ingroup taxa (the organisms of interest) and outgroup taxa (organisms outside the group of interest) for phylogenetic analysis can have a significant influence on the strength of the resulting evolutionary hypothesis. Older phylogenies often contained small numbers of ingroup species, between six and twelve, that were a mixture of very closely-related taxa and a few that were very divergent. This poor taxon sampling for this very large and diverse genus, coupled with the use of very distantly-related species as outgroups, probably lead to some spurious results and the conflicting conclusions found in the literature at the time.

In addition to problems of taxon sampling, for years the study of malaria systematics was also stymied by inherent problems in the loci chosen for analysis. Small subunit (SSU) rRNA and circumsporozoite protein (CSP) have been the workhorses of Plasmodium phylogenetics (Waters et al., 1991; Waters et al., 1993; Escalate and Ayala, 1994; Escalante et al., 1995; McCutchan, et al. 1996; Qari et al., 1996; Escalante et al., 1996; Escalante et al., 1997). Recent research indicates, however, that neither of these loci may be appropriate for evolutionary studies of Plasmodium species. SSU rRNA is a standard locus used in high-level molecular systematics, but it was later found that Plasmodium species possess separate genes of rRNA, each expressed at a different point in the life cycle, that are not evolving in a concerted manner and may still be exchanging genetic information with each other (Corredor and Enea, 1993). Several of the older phylogenies may have included a mixture of paralogs (duplicate genes) and orthologs (proper homologs), and only orthologs yield reliable gene trees. Although life-stage-specific primers have been developed for the SSU rRNA loci in Plasmodium, as gene conversion among the non-homogenized paralogs cannot be ruled out, this locus should be used with great caution. Indeed, even very recent phylogenies (Leclerc et al., 2004) using these sequences show results that cannot be reconciled with those of other loci in the genome.

Likewise, the locus CSP, which has been frequently used for evolutionary studies, may be problematic as the gene codes for a surface protein and is under strong selective pressure from the vertebrate immune system. Selectively driven, non-neutral changes in the gene may either obscure the phylogenetic signal or lead to incorrect phylogenetic inferences (Hughes and Hughes, 1995). CSP was a favored locus for study as large amounts of sequence data was available early on as the protein is of great interest to malaria science as a possible vaccine target. Improved methods for gene sequencing have allowed for better loci to be developed for evolutionary studies and CSP is now rarely used in systematics.

At this time, new loci are being developed for molecular studies, such as the mitochondrial gene for cytochrome b (Escalante et al., 1998; Perkins and Schall 2002) and the gene for the housekeeping enzyme adenylosuccinate lyase (Kedseierki et al., 2002). One caveat is that mitochondrial cytochrome b is the target of some antimalarial drugs, and mutations in the gene are known to be associated with resistance (see Vaidya, et al., 1993). Studies of these and other suitably chosen genes will hopefully lead to Plasmodium phylogenies based on the combined data from multiple loci. It should be noted, however, that the chance of identifying true neutrally evolving loci in Plasmodium species is compromised by the possibility of unrecognized selection pressure from the host immune system or (in the case of human parasites) drugs, and hence an extra measure of caution is called for.

One intuitive assumption that was challenged early on by molecular studies was the idea that human malaria parasites are closely related. Recent work now shows that, although primate parasites (with the exception of P. falciparum and P. reichenowi) grouped together, those parasites with a human host were not closely related within this group. The relatively distant relationships among human pathogens indicates that malaria most likely has an ancient association with primates (Escalante et al., 1998; Perkins and Schall, 2002; Leclerc et al., 2004, Cornejo and Escalante, 2006).

One of the most intriguing issues in malaria evolution is the mysterious origin of P. falciparum, and few issues have gone through as many significant revisions in the past 15 years. Older textbooks, and even some current ones, refer to P. falciparum as arising from a recent horizontal transfer from birds, and it is to this recent host shift that the high virulence of P. falciparum is often attributed (White, 2003). Newer research, however, refutes this hypothesis, as P. falciparum falls within a clade of mammalian parasites—distinct from and outside the group affecting birds and lizards. On the other hand, the species is also outside of the primate or rodent malarias. Rather, P. falciparum is in a divergent clade that includes only itself and its close sister species, the chimpanzee parasite, P. reichenowi.

The avian-origin hypothesis arose from some of the early molecular phylogenies that had problems due to insufficient taxon sampling and choice of genes (Waters et al., 1991; Waters et al., 1993; Escalante et al., 1995; Escalante et al., 1996; Escalante et al., 1997). First in 1996 (Qari et al., 1996) and then again in 2002 (Perkins and Schall, 2002), new evidence came to light refuting an avian-origin. Perkins and Schall (2002) published a relatively strong phylogeny based on cytochrome b with a large sample size of vertebrate parasites and an outgroup from a sister family. This cytochrome b phylogeny placed P. falciparum within the clade of mammalian parasites. In addition to the phylogenetics, this analysis included an explicit test of the avian-origin hypothesis using the Shimodaira-Hasegawa test, a phylogenetic tree-based method that calculates the likelihood of alternate trees, and rejected it.

At this time, the evidence (Qari et al., 1996; Escalate and Ayala, 1997; Perkins and Schall, 2002; Leclerc et al., 2004; Martinsen et al., 2008) indicates that the only species closely related to P. falciparum is P. reichenowi, and the two likely diverged from each other between 5 and 8 million years ago based on fossil dates of the human-chimpanzee split (Escalante and Ayala, 1994; Escalante et al., 1995).

Future phylogenetic analyses that include parasites that infect bats (several genera of malaria parasites have been described from these hosts alone) and ungulates may help to identify close relatives of P. falciparum and better understand the evolutionary history of these parasites.

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Wikipedia

List of Plasmodium species

The genus Plasmodium is a member of the order Haemosporidia. It is the largest genus within this order and currently consists of over 250 species.

The species in this genus are entirely parasitic with part of their life cycle spent in a vertebrate host and another in an invertebrate host - usually a mosquito. Vertebrates infected by members of this genus include mammals, birds and reptiles.

Host range among the mammalian orders is non uniform. At least 29 species infect non human primates; rodents outside the tropical parts of Africa are rarely affected; a few species are known to infect bats, porcupines and squirrels; carnivores, insectivores and marsupials are not known to act as hosts.

The listing of host species among the reptiles has rarely been attempted. Ayala in 1978 listed 156 published accounts on 54 valid species and subspecies between 1909 and 1975.[1] The regional breakdown was Africa: 30 reports on 9 species; Australia, Asia & Oceania: 12 reports on 6 species and 2 subspecies; Americas: 116 reports on 37 species.

Diagnostic criteria of the order Haemosporida[edit]

Currently there are ~550 species recognised in this order organised into 17 genera.[2]

The diagnostic criteria of this family are:

Diagnostic criteria of the genus Plasmodium[edit]

Note[edit]

Mammalian erythrocytes do not possess a nucleus. Although it has been suggested that the nucleus was lost in the erythrocytes better to enable them to traverse capilaries evidence for this is lacking. It appears that this loss along with the mitochondria that the erythrocytes also lose may protect the erythrocytes against oxidative stress.[3]

Subgenera[edit]

The full taxonomic name of a species includes the subgenus but this is often omitted in practice. The full name indicates some features of the morphology and type of host species. Sixteen subgenera are currently recognised.

The avian species were discovered soon after the description of P. falciparum and a variety of generic names were created. These were subsequently placed into the genus Plasmodium although some workers continued to use the genera Laverinia and Proteosoma for P. falciparum and the avian species respectively.

The 5th and 6th Congresses of Malaria held at Istanbul (1953) and Lisbon (1958) respectively recommended the creation and use of subgenera in this genus. Laverinia was applied to the species infecting humans and Haemamoeba to those infecting lizards and birds. This proposal was not universally accepted. Bray in 1955 proposed a definition for the subgenus Plasmodium and a second for the subgenus Laverinia in 1958. Garnham described a third subgenus - Vinckeia - in 1964. Several additional subgenera have been created since. The currently recognised subgenera are listed below.

Asiamoeba Telford 1988
Bennettinia Valkiūnas 1997[4]
Carinamoeba Garnham 1966
Giovannolaia Corradetti, Garnham & Laird 1963[5]
Haemamoeba Grassi & Feletti 1890
Huffia Garnham & Laird 1963
Lacertaemoba Telford 1988
Laverania Bray 1958[6]
Novyella Corradetti, Garnham & Laird 1963
Nyssorhynchus Poinar 2005
Ophidiella Garnham 1966
Papernaia Landau et al 2010[7]
Paraplasmodium Telford 1988
Plasmodium Bray 1963 emend. Garnham 1964
Sauramoeba Garnham 1966
Vinckeia Garnham 1964

Classification criteria for subgenera[edit]

The current classification scheme was developed prior to the widespread use of DNA sequence based taxonomy and is based on host and morphological criteria. Plasmodium has since been shown to be paraphytic with the genera Haemoproteus and Hepatocystis (vide infra).[8] Revision of this genus will be undertaken once sufficient DNA sequence material is available.

This forthcoming reclassification project is not unique to this genus as DNA based taxonomy is revising many traditional groupings of protozoa.

Species with mammalian hosts[edit]

Laverania

Species in this subgenus infect higher primates (including man) and have characteristic sickle shaped female gametocytes.

The type species is Plasmodium falciparum.

Plasmodium

Species infecting higher primates other than those in the subgenus Laverania are placed in the subgenus Plasmodium.

The type species is Plasmodium malariae.

Vinckeia

Parasites infecting other mammals including lower primates (lemurs and others) are classified in the subgenus Vinckeia.

The type species is Plasmodium bubalis.

Species with avian hosts[edit]

Bennettinia

Schizonts contain scant cytoplasm, are often round, do not exceed the size of the host nucleus and stick to it. Gametocytes, while varying in shape tend to be round or oval, do not exceed the size of the nucleus and stick to it.

The type species is Plasmodium juxtanucleare.

Giovannolaia

Schizonts contain plentiful cytoplasm, are larger than the host cell nucleus and frequently displace it. They are found only in mature erythrocytes. Gametocytes are elongated. Exoerythrocytic schizogony occurs in the mononuclear phagocyte system.

The type species is Plasmodium circumflexum.

Haemamoeba

Mature schizonts are larger than the host cell nucleus and commonly displace it. Gametocytes are large, round, oval or irregular in shape and are substantially larger than the host nucleus.

The type species is Plasmodium relictum.

Huffia

Mature schizonts, while varying in shape and size, contain plentiful cytoplasm and are commonly found in immature erythryocytes. Gametocytes are elongated.

The type species is Plasmodium elongatum.

Novyella

Mature schizonts are either smaller than or only slightly larger than the host nucleus. They contain scanty cytoplasm. Gametocytes are elongated. Sexual stages in this subgenus resemble those of Haemoproteus. Exoerythrocytic schizogony occurs in the mononuclear phagocyte system

The type species is Plasmodium vaughani.

Papernaia

The gametocytes are elongated. The schizonts apically or lateroapically placed and are rounded or irregularly shaped. The host nucleus may be tilted.

The type species is Plasmodium polare

Species with reptilian hosts[edit]

Although over 3200 species of lizard have been identified as hosts to Plasmodium species, only 29 species of snakes have been. All snake infecting species are placed into the sungenus Ophidiella.

Asiamoeba

The schizonts and gametocytes are greatly disparate in size (4 to 15 times).

Carinamoeba

The schizonts are small and give rise to 8 or fewer merozoites. The gametocytes like the schizonts are small.

The type species is Plasmodium minasense.

Lacertaemoba

The schizonts are medium-sized and undergo 3 to 5 nuclear divisions. The gametocytes are medium-sized

Paraplasmodium

The schizonts are of medium size. Exoerythrocytic schizonts may be produced in both fixed and wandering host cells. The gametocytes are large. One species in this sub-genus is capable of merogony in a vector of the Lutzomyia genus.

Sauramoeba

Large schizonts giving rise to 12 or more merozoites. The gametocytes like the schizonts are large. The asexual stages tend to disappear from the lymphocytes once the gametocytes appear in the lymphocytes.

The type species is Plasmodium agamae.

Ophidiella

The species in this subgenus infect only snakes.

The type species is Plasmodium weyoni.

Species with unknown hosts[edit]

One species has been identified from Dominican amber - Plasmodium dominicum. The vertebrate host of this species is unknown but it seems likely that it may have been a bird.

Nyssorhynchus

The type species is Plasmodium dominicum.

Phylogenetics[edit]

Although the evolution of this genus has been studied by a number of authors, details are still being elucidated. A brief summary of the pattern that has emerged is as follows:

The most basal split in the genus is between the reptile/bird species and the mammalian species. The bird/reptile clade appears to be related to the genera Haemoproteus, Leukocytozoon and Polychromophilus. The genus Hepatocystis appears to have evolved from with the mammalian species clade. Within the mammalian species the subgenus Laverinia appears to be basal with the subgenus Plasmodium and the rodent species being sister clades. Hepatocystis appears to have diverged after the separation of the rodent species. The species infecting lemurs may belong in the subgenus Plasmodium instead of their current placement within the subgenus Vinckeia.

Within the subgenus Plasmodium, P. vivax groups with an Asian clade which appears to be rooted in Africa. P. malaria and P. ovale both belong to an African clade and are more closely related to each other than to P. vivax. Within the subgenus Laverinia P. falciparum and P. reichenowi form a clade while the other four known species form a second clade.

There are a number of additional species in these taxa that await full description so changes to the branching order are likely. However the overall arrangement outlined above seems to be supported by a number of studies by different authors and is unlikely to change. Given the recently recognised paraphytic nature of several of the taxa above, the introduction of new genera and possibly families in the near future seems highly likely.

Relations with other Haemosporidian genera[edit]

While most phylogenetic trees have tended to agree that Plasmodium has descended from Leukocystis or Haemoproteus like species a Bayesian phylogenetic reconstruction suggests that Plasmodium may be the ancestral genus that has given rise to Haemoproteus and other genera.[9] Further study in this area is required.

Another Bayesian analysis has suggested the following taxonomy: Mammalian Plasmodium and Hepatocystis are sister clades with Hepatocystis having evolved from within the genus Plasmodium; the bird and reptile species are intermixed and basal to the mammalian Plasmodium/Hepatocystis species; the reptilan/bird Plasmodium species are a sister clade to the genus Polychromophilus; Leukocytozoon and Haemoproteus and sister clades; the Leukocytozoon/Haemoproteus clade is a sister to the Parahaemoproteus clade; and the Parahaemoproteus/Haemoproteus/Leukocytozoon clade is a sister to the reptilan/bird Plasmodium/Polychromophilus clade.[9] This grouping is supported by previous results.[10]

A study of DNA sequences suggests that the genus is paraphytic with Hepatocystis being related to the mammalian species and Polychromophilus being related to the reptile species.[11] This study also supports the ancestor of Plasmodium being a Leucocytozoon like species and that Plasmodium is more closely related to the Haemoproteus - specifically the subgenus Parahaemoproteus - than to Leucocytozoon.

A paper by Blanquart and Gascuel[12] examined Plasmodium 84 mitochondial sequences and included Hepatocystis, Haemoproteus and Leukocytozoon sequences. The results agree with the previous analyses showing that Hepatocystis, Haemoproteus and Plasmodium appear to be derived from a Leukocytozoon ancestor. Hepatocystis appears to be a sister group to the great ape-rodent clade with the lower primate clade being ancestral to all three. In terms of Plasmodium subgenera they suggest that the subgenus Plasmodium is ancestral to both Laverania and Vinckeia.

A study of parasites infecting bats found that the bats were infected by species of the genera Hepatocystis, Plasmodium, Polychomophilus and Nycteria.[2] A phylogenetic tree which included these genera along with Haemoproteus and Leukocytozoon species was examined. As before Leukocytozoon was basal in this tree. The next clade to diverge was that of the Haemoproteus species. The remaining genera lay within the currently established genus Plasmodium.

Within this genus the first to diverge were the avian and reptile species. The next clade to diverge was that of the Polychomophilus species. This was followed in branching order by the Nycteria species. The subgenus Laverinia was the next to diverge followed by the subgenus Vinckeia. The crown of the tree was formed by the subgenus Plasmodium and the genus Hepatocytis. This tree did not support the inclusion of P. ovale in the subgenus Vinckeia but agreed with previous analyses suggesting that P. malaria is more closely related to the Asian clade than P. ovale is. Several of the bat infecting Plasmodium species appear to be related to the rodent species.

Bats appear to have evolved ~66 million years ago in Africa[13] which - assuming that the phylogenetic tree in Schaer et al is correct - places an upper limit on the date for the evolution of the mammalian species of Plasmodium.

Relations with non Haemosporidian genera[edit]

The Piroplasma are usually considered to be the closest relations to the Haemosporidians. Based on the evolution of their vectors (ticks) may have evolved ~300 million years ago.[14] The vectors of Babesia and Theileria - ticks - evolved 350 million years ago ± 23 million years ago.[15] The hard (Ixodidae) and soft bodied (Argasidae) ticks diverged 290 million years ago± 23 million years ago. The most likely place of origin of the ticks is Northern Gondwana and most probably within the region that now constitutes Eastern Africa.

A molecular Bayesian study of Babesia and Theileria species along with Plasmodium species suggests that Babesia and Theileria are sister clades and that they diverged from Plasmodium ~56.5 million years ago (95% credible interval: 86.9 million years ago - 28.2 million years ago)[16] The dating in this study used a date of 12.5 million years ago for the origin of the genus Plasmodium.[17] The authors also estimated that Theileria evolved 23.38 million years ago (95% crdibile interval 11.1 million years ago36.7 million years ago) and that Babesia evolved 25.7 million years ago (95% credible interval 12.8 million years ago40.7 million years ago)

Another analysis suggests that Babesia and Theileria are more closely related to the adeleid species than to Plasmodium.[18]

An examination of sequences from Babesiidae, Cryptosporiidae, Eimeriidae, Plasmodiidae, Sarcocystiidae, Theileriidae, a Perkinsus species and 2 dinoflagellates suggests that Plasmodium and Cryptosporidium are sister taxa and that Hepatozoon is basal to them.[19]

Morrison has shown using molecular data that the Haemosporidia are nested within the gregarines and that this clade is distinct from the piroplasms.[20] This latter clade is a sister group of the coccidians.

Examination of the actin genes suggests that Plasmodium is more closely related to the coccidians than to the Babesia/Theileria clade.[21] It also suggests that Cryptosporium is basal in the Apicomplexa: this latter finding is consistent with other analyses.

Phylogenetic trees[edit]

A number of useful phylogenetic trees of this genus have been published:

Tree of Life website
American Museum of Natural History
PLOS site
Paper on Plasmodium
Paper on Plasmodium
Paper on Plasmodium
Paper on Plasmodium

From these trees it is clear that:

  • The trees are consistent with the origin of Plasmodium from Leukocytozoon
  • The genus Hepatocystis is nested within (paraphytic with) the genus Plasmodium and appears to lie within the primate-rodent clade[22]
  • The rodent and primate groups are relatively closely related
  • The primate (subgenus Plasmodium) and rodent species (subgenus Vinckeia) form distinct groups
  • P. falciparum and P. reichenowi (subgenus Laverania) branched off early in the evolution of this genus
  • The 'African' (P. malaria and P. ovale) and 'Asian' (P.cynomogli, P. semiovale and P. simium) species tend to cluster together into separate clades. Interestingly P. gonderi - a species isolated in Africa - groups with the Asian clade.
  • P. vivax clusters with the 'Asian' species.
  • The rodent species (P. bergei, P. chabaudi and P. yoelli) form a separate clade.
  • The species infecting humans do not form a single clade.[23]
  • The genus Haemoproteus appears to lie within the bird-lizard clade
  • The lizard and bird species are intermingled
  • Although Plasmodium gallinaceum (subgenus Haemamoeba) and Plasmodium elongatum (subgenus Huffia) appear be related here so few bird species (three) have been included, this tree may not accurately reflect their real relationship.
  • The bird species (P. juxtanucleare, P. gallinaceum and P. relictum) form a clade that is related to the included Leukocytozoon and Haemoproteus species.
  • While no snake parasites have been included these are likely to group with the lizard-bird division
  • Hepatocystis seems to lie within Plasmodium and may be related to the primate clade

The bird and lizard species are intermixed as previously found.

An analysis of the rodent genera (Plasmodium berghei, Plasmodium chabaudi, Plasmodium vinckei and Plasmodium yoelii) suggests that that these species may actually be species complexes.[24] The separation of P. chabaudi and P. vinckei has been estimated to be between 3 million years ago and 13 million years ago while that of P. berghei and P. yoelii has been placed at 1 million years ago and 6 million years ago.

A paper that included five unnamed lemur species suggested that P. ovale is more closely related to the lemur species than to the other primate ones.[25] It also suggested that the lemur/P. ovale clade is a sister clade of the rodent species. While this is consistent with the placement of the lemur and rodent species in the subgenus Vinckeia it is inconsistent with the current placement of P. ovale within the subgenus Plasmodium. This paper also supports a basal divergence within the mammalian species into the subgenus Laverinia and the others. The subgenera Plasmodium and Vinckeia with the exception of P. ovale appear to be sister clades.

Analysis of the apicoplast genes auggests that P. ovale is related to the rodent species.[26] This is consistent with its relationship with the lemur species in the subgenus Vinckeia.

Other analyses[edit]

Examination of the protease gene (SERA) in 18 species[27] has shown that the ancestral state had only a single gene and that gene duplications have occurred in the extant species. This paper confirms the groupings found elsewhere with an Asian clade. The rodent species seem to be more closely related to the Laverania subgenus than does the subgenus Plasmodium.

A deletion mutation of ~100 base pairs including part of the LS1 rRNA gene is found in the sequences of two African species - P. gonderi and an undescribed parasite taken from a mandrill - and 2 Asian species - P. cynomolgi and P. simiovale.[28] This mutation was not found in the other species examined (Leucocytozoon caulleryi, Leucocytozoon sabrazesi, P. bergei, P. chabaudi, P. falciparum, P. floridense, P. gallacium, P. fragile, P. juxtanucleare, P. knowelsi, P. mexicanum, P. reichenowi, P. relictum, P. simiae, P. vivax, P. yoelii and two unnamed Haemoproteus species.) These mutations are rare events and strongly suggests these species are related.

Another paper suggests that after the mammalian-reptile/bird species split that the subgenus Laverina is basal among the mammal species.[29] This study did not include mammalian infecting species other than primate and rodent species and for this reason Laverina may not be as basal as the study suggests. The remaining branching order is consistent with other analyses placing the rodent species as the first branch after the P. falciparum/P. reichenowi clade. It places P. malaria and P. ovale as being more closely related to each other than to P. vivax. This is consistent with the proposed Asian origin of P. vivax.

Although bird malaria species use a variety of mosquito vectors from the genera Aedes, Anopheles, Culex, Culiseta, Mansonia and Psorophora, all mammalian species use vectors only from the genus Anopheles.[30] This host switch seems to have been associated with a specilization with a particular genus of mosquito.

The ability to store haemozoin appears to have evolved only once in the common ancestor of Haemoproteus, Hepatocystis and Plasmodium.[30]

A study of the relationships between Haemocystis, Haemoproteus, Leucocytozoon and Plasmodium suggests that (1) Leucocytozoon is basal (2) Haemoproteus is a sister clade to the remainder (3) Parahaemoproteus is a sister to Plasmodium and (4) Haemocystis is nested within Plasmodium.[30] As before the bird/lizard species form a distinct clade.

In birds the Haemoproteus and Leucocytozoon species rarely change transmission area.[31] These parasites are restricted to one resident bird fauna over a long evolutionary time span and are not freely spread between the continents with the help of migratory birds. Lineages of the genus Plasmodium in contrast seem more freely spread between the continents. This suggests that the origin on the genus Plasmodium may have coincided with the ability to transfer between avian hosts more easily than the other genera.

Molecular clock estimates[edit]

All dates estimated so far using a molecular clock should probably be regarded with some suspicion given the existing disagreements between the various authors.

The branching order suggested by other analyses concurs with an analysis of the mitochondial genes[32] This latter paper puts the divergence between the reptile-bird and mammal clades at 38.4 million years ago ± 3.2 million years ago (Mya). Other divergence times reported include

An estimate of the dates of evolution of several species[33] using the date of separation of the African species P. gonderi and the Asian clade at 10 million years ago gives estimates as follows:

Analysis of 45 single copy nuclear genes from eight species (P. berghei, P. chabaudi, P. falciparum, P. gallinaceum, P. knowlesi, P. reichenowi, P. vivax, P. yoelii) using several different phylogenetic methods suggest a divergence date between Theileria and Plasmodium between 294 million years ago and 314 million years ago.[34] Estimates of the mutation rates suggest a date of divergence between P. falciparum and P. reichenowi between 5 million years ago and 7 million years ago.

The estimated date of divergence between P. vivax and P. knowlesi was between 15 million years ago and 46 million years ago. This latter period coincides with the radiation of the Old World monkeys which these parasites infect. The date of divergences between P. berghei, P. chabaudi and P. yoelii was estimated to be between 34 million years ago and 25 million years ago. The main radiation of the rodent family Muridae occurred ~24 million years ago.

A paper based on the analysis of 22 nuclear genes suggests a radiation of malarial parasites within the Oligocene (34-23 million years ago).[35]

Another paper[32] examining the dates of evolution using the concatenated sequences of the cytochrome c oxidase III, cytochrome c oxidase I and cytochrome b genes - all from the mitochondrion - suggested the following dates for the evolution of the species examined (P. coatneyi, P. cynomolgi, P. falciparum, P. fieldi, P. fragile, P. gonderi, P. hylobati, P. inui, P. knowlesi, P. malariae, P. ovale, P. reichenowi, P. simiovale, P. vivax) was as follows:

Asian-African primate clade divergence: 12 million years ago-19 million years ago

Primate-rodent clade divergence: 15 million years ago-30 million years ago

Reptile/bird-mammal clade divergence: 20 million years ago-30 million years ago

An estimation of the date of evolution of this genus based upon the mutation rate in the cytochrome b gene places the evolution of P. falciparum at 2.5 million years ago.[17] The authors also estimated that the mammalian species of this genus evolved 12.8 million years ago and that the order Haemosporida evolved 16.2 million years ago. While the date of evolution of P. falciparum is consistent with alternative methods, the other two dates are considerably more recent than other published estimates and probably should be treated with caution.

Another paper which examined primate, rodent, lemur, bird and reptile species suggests that the genus originated between 30 million years ago and 50 million years ago.[25] The split between the reptile/bird and mammalian species occurred between 31.4 million years ago and 47.6 million years ago. The first division in the mammalian species was between Laverinia and the others species. The separation of P. falciparum and P. reichenowi was estimated to be between 3.6 million years ago and 7.9 million years ago. The bonobo strains of P. falciparum of were the closest relations to those of humans. This analysis grouped P. ovale with the lemur species and this clade as a sister clade to the rodent species. While this is consistent with the current placement of the lemur species with the rodent species in the subgenus Vinckeia, it is inconsistent with the current placement of P. ovale in the subgenus Plasmodium. The date of separation of P. ovale from the lemur species was estimated to be 25 million years ago and 35 million years ago and their date of divergence from the rodent species was dated to between 30 million years ago and 50 million years ago. The rodent species first diverged between 10 million years ago and 20 million years ago. P. atherui appears to be more closely related to the P. berghei/P. yoelli clade than to P. chabaudi. P. malariae evolved between 20 million years ago and 30 million years ago and is more closely related to P. vivax than to P. ovale. P vivax and P. cynomogli last shared an ancestor between 2.2 million years ago and 4.5 million years ago. The origin of the Asian clade was placed between 5 million years ago and 8.2 million years ago.

Laverania[edit]

Four species (P. billbrayi, P. billcollinsi, P. falciparum and P. reichenowi) form a clade within the subgenus Lavernia. This subgenus is more closely related to the other primate species than to the bird species or the included Leuocytozoon species. Both P. billbrayi and P. billcollinsi infect both the chimpanzee subspecies included in this study (Pan troglodytes troglodytes and Pan troglodytes schweinfurthii). P. falciparum infects the bonbo (Pan paniscus) and P. reichenowi infects only one subspecies (Pan troglodytes troglodytes). Caution has been raised about the adequacy of the description of these new species.[36]

A report of a new species that clusters with P. falciparum and P. reichenowi in chimpanzees has been published, although to date the species has been identified only from the sequence of its mitochondrion.[37] Further work will be needed to describe this new species, however, it appears to have diverged from the P. falciparum- P. reichenowi clade about 21 million years ago. A second report has confirmed the existence of this species in chimpanzees.[38] This report has also shown that P. falciparum is not a uniquely human parasite as had been previously believed. A third report on the epidemiology of P. falciparum has been published.[39] This study investigated two mitochondrial genes (cytB and cox1), one plastid gene (tufA), and one nuclear gene (ldh) in 12 chimpanzees and two gorillas from Cameroon and one lemur from Madagascar. Plasmodium falciparum was found in one gorilla and two chimpanzee samples. Two chimpanzee samples tested positive for Plasmodium ovale and one for Plasmodium malariae. Additionally one chimpanzee sample showed the presence of P. reichenowi and another P. gaboni. A new species - Plasmodium malagasi - was provisionally identified in the lemur. This species seems likely to belong to the Vinckeia subgenus but further work is required.

A study of ~3000 wild ape specimens collected from Central Africa has shown that Plasmodium infection is common and is usually with multiple species.[40] The ape species included in the study were western gorillas (Gorilla gorilla), eastern gorillas (Gorilla beringei), bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). 99% of the strains fell into six species within the subgenus Laverina. P. falciparum formed a monophyletic lineage within the gorilla parasite radiation suggesting an origin in gorrilas rather than chimpanzees.

It has been shown that P. falciparum forms a clade with the species P. reichenowi.[41] This clade may have originated between 3 million and 10000 years ago. It is proposed that the origin of P. falciparum may have occurred when its precursors developed the ability to bind to sialic acid Neu5Ac possibly via erythrocyte binding protein 175. Humans lost the ability to make the sialic acid Neu5Gc from its precursor Neu5Ac several million years ago and this may have protected them against infection with P. reichenowi.

Another paper has suggested that the P. falciparum isolates found in apes are derived from humans and that P. falciparum and P. reichenowi diverged when humans and chimpanzees/gorillas did (between 5 million years ago and 7 million years ago million years ago).[42]

It is considered that P. falciparum in humans originated from a single transmission event and that the great apes do not represent a potential reservoir for on going transmission.[43]

The origin of P. falciparum in humans seems likely to have been from bonobos rather than gorillas or chimpanzees.[29]

Another estimate of the most recent common ancestor of the extant strains that has been published is 452,000 years ago.[44]

A review of this subgenus has been published[45] Based on the analysis of the cytochrome b gene the relationships in this subgenus appear to as follows: P. falciparum and P. reichenowi are sister species. Their closest relation is P. billcollinsi. P. gaboni and P. billbrayi are sister species whose closest relation is P. gora. P. gorb is more closely related to the P. falciparum/reichenowi/billcollinsi clade than the P. gaboni/billbrayi/gora clade. This putative taxonomy will need confirmation from other DNA studies. A second study seems to confirm this proposed grouping.[46]

The dates of the evolution of the species within the subgenus Laverania have been estimated as follows:[29]

Another estimate using the mutation rate (1.2 x 10−8 subsititutions/site/year) of the cytochrome b gene placed the spread of P. falciparum to humans at 365,000 years ago (95% credible interval: 112,000 to 1,036,000 years).[47]

Revised names have been proposed for the P. gora and P. gorb species - Plasmodium blacklocki and Plasmodium adleri respectively.[48] These names were chosen to honour the malariologists Saul Adler (1895–1966) and Donald Blacklock (1879–1953). It has also been proposed that the P. falciparum strains infecting gorillas should be renamed Plasmodium praefalciparum. This proposal appears to have been accepted.[46] The species P. billbrayi seems to be synonymous with earlier named P. gaboni.

Host-parasite relations:

  • P. falciparum has been isolated from chimpanzees, gorillas and humans. The non human strains may be reclassified as P. praefalciparum.
  • P. reichenowi has been isolated from chimpanzees.
  • P. billcollinsi has been isolated from chimpanzees.
  • P. billbrayi has been isolated from chimpanzees.
  • P. gaboni has been isolated from chimpanzees.
  • P. adleri has been isolated from gorillas.
  • P. blacklocki has been isolated from gorillas.

It appears that P. falciparum has been introduced into South America on several occasions.[49] The extant strains fall into two clades - one northern and one southern. The most probable origin of these strains is Africa and it seems that they were introduced with the slave trade.

Analysis of 45 single copy nuclear genes from eight species (P. berghei, P. chabaudi, P. falciparum, P. gallinaceum, P. knowlesi, P. reichenowi, P. vivax, P. yoelii) using several different phylogenetic methods suggest a divergence data between 294 and 314 between Theileria and Plasmodium.[34] Estimates of the mutation rates suggest a date of divergence between P. falciparum and P. reichenowi between 5 million years ago and 7 million years ago.

Analysis of polymorphisms in the mitochondrial[50][51] genes suggests a sub Saharan origin for P. falciparum with separate colonisations of Southeast Asia and Oceania. Given the distributions of the other members of Laverinia it seems likely all the known members of this subgenus originated in Africa.

Plasmodium[edit]

Colobine and macaque monkeys migrated from Africa into the Eurasian continent 10 and 6 millions of years ago respectively and became the ancestors of the extant Asian old world monkey species.[52] Asian old world monkey malaria parasite species infect both colobine and macaque monkeys. The existing divergence between the Asian and African clade of this subgenus seems likely to have been caused by intercontinental allopatric speciation along with that of their hosts.

Malaria parasites of the lemurs are not traditionally grouped with the subgenus Plasmodium being placed rather within subgenus Vinckeia. This classification may not be correct.[53] Based on an analysis of the mitocondria, these parasites seem to group with the others infecting primates. The origin of the primate infecting species (excluding those in the Laverina subgenus) may date back to the Eocene - a time when the primate radiation began. This analysis also suggests that the species infecting gorillas and humans may have originated in chimps.

Plasmodium: Asian clade[edit]

At least nine species belong to the 'Asian' clade of Plasmodium. These species include Plasmodium coatneyi, Plasmodium cynomolgi, Plasmodium fieldi, Plasmodium fragile, Plasmodium inui, Plasmodium hylobati, Plasmodium simiovale, Plasmodium simium and Plasmodium vivax.

As a rule (with the noticeable exception of P. knowesli), the Asian species have a 72-hour intra erythroctytic life cycle.

Analysis of the merozoite surface protein in ten species of the Asian clade suggest that this group diversified between 3 and 6.3 million years ago - a period that coincided with the radiation of the macques within South East Asia.[54] The inferred branching order differs from that found from the analysis of other genes suggesting that this phylogenetic tree may be difficult to resolve. Positive selection on this gene was also found.

In an analysis of the SSU rRNA gene it was found that all Asian simian Plasmodium species have a single S-type-like gene and several A-type-like genes.[55] A 50 residue insertion in the V7 variable region near the stem 43 is shared exclusively by the S-type like sequences of the Asian simian Plasmodium species and the S- and O-type sequences of P. vivax. This is consistent with their shared ancestry.

Plasmodium vivax may have originated in Asia and the related species Plasmodium simium appears to be derived through a transfer from the human P. vivax to New World monkey species in South America. This was proposed in a study of howler monkeys near São Paulo, Brasil.[56]

Another paper has suggested an African origin for P. vivax.[57]

Time to most recent common ancestor

P. vivax appears to have evolved between 45,000 and 82,000 years ago from a species that infects south east Asian macques.[58] This is consistent with the other evidence of a south eastern origin of this species. A second estimate put the earliest date of the evolution of P. vivax at 265,000 years.[59]

An estimate of the date of origin of P. vivax has placed it at 768,000 years ago.[44]

An estimate of the time of origin of P. vivax based on nuclear genes suggests that it originated between 232,228 to 303,030 years ago.[60] It may have appeared in India between 79,235 and 104,008 years ago.

A study of P. vivax in the Americas suggests that the strains in Venezuela and northeastern Brazil diverged from the others ~30,000 years ago.[61] This separation may have occurred before the parasite was introduced into South America.

The most recent common ancestor of the extant P. knowlesi strains has been estimated to have appeared 257,000 (95% credibility interval 98,000–478,000) years ago.[33] P. knowlesi underwent a rapid population growth between approximately 30,000 and 40,000 years ago. This era follows the growth in the human population in this area (~50,000 years ago).[62]

Branching order

P. coatneyi and P. inui appear to be closely related to P. vivax.[22]

P. vivax and P. knowesli appear to have diverged 25–30 million years ago.[42]

P. gonderi appears to be basal in this clade.[63] This is consistent with its African distribution rather than the mainly Asian distribution of the other species in this group.

Several of the 'Asian' clade - Plasmodium coatneyi, Plasmodium cynomolgi, Plasmodium fragile, Plasmodium inui, Plasmodium fieldi, Plasmodium hylobati, Plasmodium inui, Plasmodium knowlesi and Plasmodium simiovale and an African species Plasmodium gonderi - have a single S-type-like gene and several A-type-like genes. It seems likely that these species form a clade within the subgenus Plasmodium.

The 'Asian' species form a clade with P. simium and P. vivax being clearly closely related as are P. knowseli and P. coatneyi and P. fragile;[63] similarly P. brazillium and P. malariae are related. P. hylobati and P. inui are closely related. P. fragile and P. gonderi appear to be more closely related to P. vivax than to P. malariae.

An analysis of four apicoplast genome-encoded genes (small subunit rRNA, large subunit rRNA and caseinolytic protease C) of nine 'Asian' species (P. coatneyi, P. cynomolgi, P. fieldi, P. fragile, P. hylobati, P. inui, P. knowlesi, P. simiovale and P. vivax) and the African species P. gonderi suggests that P. coatneyi and P. knowlesi are closely related and that P. fragile is the species most closely related to these two.[64] Also P. vivax and P. cynomolgi appear to be related.

The pattern emerging from this data suggests that the ancestor of P. gonderi and the 'Asian' clade (P. coatneyi, P. cynomolgi, P. fieldi, P. fragile, P. hylobati, P. inui, P. knowlesi, P. simiovale and P. vivax) infected a primate host - perhaps the ancestor of the extant Rhesus monkey - and migrated with its vertebrate host from Africa to Asia via the Middle East. The Asian branch then gave rise to several clades - P. fragile-P. coatneyi/P. knowlesi, P. hylobati/P. inui and P. cynomolgi - P. simium/P. vivax. P. fieldi, P. simiovale and P. vivax appear to be relatively early diverging species within this clade.[63] P. fieldi and P. simiovale appear to be each other's closest relations.

A summary of the currently understood branching order is as follows:

  • P. gondori - Asian clade
  • P. fieldi, P. simiovale, P. vivax, P. simium, P. cynomolgi, P. inui - P. fragile, P. coatneyi, P. knowlesi, P. hylobati
  • P. vivax/P. simium - P. fieldi, P. simiovale, P. cynomolgi, P. inui
  • P. cynomolgi/P. inui - P. fieldi/P. simiovale
  • P. fragile/P. coatneyi - P. knowlesi/P. hylobati

This branching order may have to be revised as more data becomes available. The timing of these events is still rather uncertain.

The African species P. georgesi appears to be a close relation of P. gondori.

Another paper suggests that P. coatneyi and P. knowlesi are sister species while P. hylobati and P. inui are also sister species.[29] This analysis supports the grouping of P. fieldi and P. semiovale as sister species with their closest relation being P. cynomogli. It also agrees with previous analyses that place P. simium and P. vivax as sister species. It also agrees that P. gondori is the African species most closely related to the Asian clade.

This branching order may have some difficulties. A deletion of the LS1 rRNA gene of P. gonderi P. cynomolgi and P. simiovale has been reported.[28] This mutation was not found in the other species of this group that were examined - P. fragile, P. knowelsi, P. simiae and P. vivax. These mutations are rare and suggest a relationship between the first three species to the exclusion of the others.

Host relations

P. cynomolgi, P. inui and P. knowlesi infect primates of the genus Presbytis.

P. cynomolgi, P. fieldi, P. inui, P. knowlesi and P. semiovale infect primates of the genus Macaca.

P. georgesi and P. gondori infect primates of the genus Cocerebus.

P. gondori infects primates of the genus Mandillus.

Additional species

Within the 'Asian' clade are three unnamed potential species. One infects each of the two chimpanzee subspecies included in the study (Pan troglodytes troglodytes and Pan troglodytes schweinfurthii).[39] These appear to be related to the P. vivax/P. simium clade.

A new species - yet to be formally described - has been reported from orangutans (Pongo pygmaeus) in Indonesia.[25] This species was identified from mitochondrial DNA in the blood of the hosts. It appears to be related to the other members of the Asian clade.

Another as yet unnamed species likely to belong to this group has been identified in the mandrill (Mandrillus sphinx).[28]

Plasmodium: African clade[edit]

The species infecting Old World monkeys (subgenus Plasmodium) seem to form a clade.

P. ovale is more closely related to P. malariae than to P. vivax.[63]

Plasmodium ovale has recently been shown to consist of two cocirculating species - Plasmodium ovale curtisi and Plasmodium ovale wallikeri.[65] These two species can only be distinguished by genetic means and they separated between 1 million years ago and 3.5 million years ago. A second estimate has placed the separation of these species at 4.5 million years ago (95% confidence interval 0.7-7.7 Mya)[63]

P. ovale, based on an analysis of the apicoplast genome, appears to be related to the rodent species suggesting an ancestral host switch.[26]

One paper has reported a strain of malaria in a chimpanzee with a mitochondrial sequence identical to that of P. ovale and a second closely related to it.[66] It seems likely as has been proposed earlier that P. ovale may have an animal reservoir.

Two unnamed potential species infect the bonbo (Pan paniscus) and these are related to the P. malariae/P. brazillium clade.

The species P. gonderi appears to be the closest relation to the Asian clade.

Rodent species[edit]

Although the branching order among the mammalian clades has not yet been determined the branching order in the rodent infections species has been studied.[24][67] The rodent parasites (P. berghei, P. chabaudi, P. vinckei and P. yoelii) seem to form a distinct clade. P. berghei and P. yoelii appear to be sister species as do P. chabaudi and P. vinckei. The separation dates between P. berghei and P. yoelii has been estimated to be 4.5 million years ago (95% credibility interval 2.5 - 6.0); that between P. chabaudi and P. vinckei has been estimated to be 9 million years ago (95% credibility interval 5.5 - 12.6); and that between the P. berghei/P. yoelii and P. chabaudi/P. vinckei clades to be 12.5 million years ago (95% credibility interval 9.0 - 17.5). These estimates are consistent with those from another paper that included a number of primate infecting species.[32]

P. atheruri appears to be the sister species of P. vinckei.[8]

Notes[edit]

A recently (2009) described species (Plasmodium hydrochaeri) that infects capybaras (Hydrochaeris hydrochaeris) may complicate the phylogentics of this genus.[68] This species appears to be most similar to Plasmodium mexicanum a lizard parasite. Further work in this area seems indicated.

Unlike other eukaryotes studied to date Plasmodium species have two or three distinct SSU rRNA (18S rRNA) molecules encoded within the genome.[55] These have been divided into types A, S and O. Type A is expressed in the asexual stages; type S in the sexual and type O only in the oocyte. Type O is only known to occur in Plasmodium vivax at present. The reason for this gene duplication is not known but presumably reflects an adaption to the different environments the parasite lives within.

It has been reported that the C terminal domain of the RNA polymerase 2 in the primate infecting species (other than P. falciparum and probably P. reichenowei) appears to be unusual[69] suggesting that the classification of species into the subgenus Plasmodium may have an evolutionary and biological basis.

It is known from many written historical sources that P. vivax malaria was endemic in the wetlands of England from the 1500s until the 20th century.[70] It is suspected that this disease was introduced by the Romans sometime before 400 AD. It seems likely that it remained endemic in these areas at least up to 1000 AD.

A study in Senegal of 25 strains isolated there suggests that P. falciparum underwent a major (60-fold) population expansion of ~20,000-40,000 years ago.[71]

A population study based on isolates from several countries suggests that distinct clustering of continental populations - Africa, Southeast Asia and Oceania - has occurred.[72] Within these grouping there has been some further clustering - West Africa versus East Africa, Thailand versus Cambodia. No distinction was identified between isolates from Mali and Burkina Faso.

Host range[edit]

Because of the number of species parasited by Plasmodium further discussion has been broken down into following pages:

Criteria used for speciation[edit]

The vertebrate host is the first criterion used for speciation and may be sufficient alone to determine the subgenus as in Ophidiella and Vinckeia. The morphological features of the parasite itself most commonly used to describe a species include the number of pigment granules, the degree of encirclement of the host nucleus, the size of the parasite, the degree of host nucleus displacement and the degree of host cell enlargement.

List of species[edit]

Unnamed species[edit]

At least one species has been isolated from the mandrill (Mandrillus leucophaeus) that awaits full publication. It is currently known as Plasmodium sp. DAJ-2004.

At least one species related to P. ovale appears to be present in chimpanzees. It is known only from a DNA sequence and awaits description.

P. vivax strains can be separated into two distinct types depending on the organisation of the A and S rRNA genes.[73] A gene conversion occurred in an Old World strain and this mutated strain give rise to a new calde of parasites in the New World. The Old World strains were subsequently re introduced - possibly via the slave trade - and these are related to the monkey parasite P. simium. The specific name Plasmodium collinsi has been proposed for the New World strains but this has not yet been accepted.

A second mutation is present in the ORF 470 gene of the plasmid in the New World P. vivax strains. This protein is highly conserved. In the Old World strains of P. vivax and its relations a valine is present. In the New World strains this residue has been replaced by an isoleucine (G -> A in the first codon position).

Two separate strains of P. vivax can be identified on the basis of the circumsporozoite protein (CSP) gene.[74] Both of these alleles can be found in P. simium and they occur both in the New and Old Worlds. This suggests a complex history of transmission across the world and between species.

Another as yet unnamed species was isolated from humans in Madang, Papua New Guinea in 1993.[75] This species differed immunologically and genetically from then generally recognised species infecting humans. Additional isolates of this putative species were also found in Sepik also in Papua New Guinea, Brazil, Indonesia and Madagascar.[76] The circumsporozoite protein of this species appears to be identical to that of Plasmodium semiovale. At least two species of mosquito Anopheles deaneorum and Anopheles oswaldoi appear to be capable of transmitting this parasite.[77] These reports have not gone unchallenged and the status of this putative species is unclear at present.[78]

Species grouped by subgenus[edit]

This listing while currently incomplete will be updated when the relevant information becomes available.

Asiamoeba
Bennetinia
Carinamoeba
Giovannolaia
Haemamoeba
Huffia
Lacertamoeba
Laverania
Novyella
Nyssorhynchus
Ophidiella
Papernaia
Paraplasmodium
Plasmodium
Sauramoeba
Vinckeia

Species subsequently reclassified into other genera[edit]

The literature is replete with species initially classified as Plasmodium that have been subsequently reclassified. With the increasing use of DNA taxonomy some of these may be once again be classified as Plasmodium. This appears increasing likely as it has been shown that Hepatocystis and Polychromophilus appear to lie within the genus Plasmodium.

The following species have been classified into the genus Hepatocystis:

The following species have been classified into the genus Haemoemba:

The following species has been classified into the genus Garnia:

The following species has been classified into the genus Fallisia:

The following species has been classified into the genus Polychromophilus:

Species now considered to be junior synonyms[edit]

P. osmaniae and P. shortii are currently considered to be junior synonyms of P. inui.

Species of dubious validity[edit]

The following species that have been described in the literature are currently regarded as being of questionable validity (nomen dubium).

References[edit]

  1. ^ Ayala S.C. (1978). "Checklist, host index, and annotated bibliography of Plasmodium from reptiles". J. Eukaryot. Microbiol. 25 (1): 87–100. doi:10.1111/j.1550-7408.1978.tb03874.x. 
  2. ^ a b Schaer J, Perkins SL, Decher J, Leendertz FH, Fahr J, Weber N, Matuschewski K (2013) High diversity of West African bat malaria parasites and a tight link with rodent Plasmodium taxa. Proc Natl Acad Sci USA
  3. ^ Zhang ZW, Cheng J, Xu F, Chen YE, Du JB, Yuan M, Zhu F, Xu XC, Yuan S (2011) Red blood cell extrudes nucleus and mitochondria against oxidative stress. IUBMB Life 63(7):560-565. doi:10.1002/iub.490.
  4. ^ Valkiunas G. (1997). Bird Haemosporidia. Institute of Ecology, Vilnius
  5. ^ Corradetti, A.; Garnham, P. C. C.; Laird, M. (1963). "New classification of the avian malaria parasites". Parassitologia 5: 1–4. 
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Plasmodium species infecting birds

Species in six subgenera of Plasmodium infect birds - Bennettinia, Giovannolaia, Haemamoeba, Huffia, Novyella and Papernaia.[1] Giovannolaia appears to be a polyphytic group and may be sudivided in the future.[2]

Avian host records[edit]

  • P. gundersi - eastern screech owl (Otus asio)
  • P. paranucleophilum - South American tanager
  • P. parvulum - vanga species

Subspecies of avian malaria[edit]

  • P. nucleophilum has at least one subspecies - P. nucleophilum toucani
  • P. relictum has been divided into subspecies: P. relictum capistranoae, P. relicturn matutinum, P. relictum quentini and P. relictum relictum.

Interrelatedness

  • P. durae is related to P. asanum, P. circumflexum, P. fallax, P. formosanum, P. gabaldoni, P. hegneri, P. lophrae, P. lophrae, P. pediocetti, P. pinotti, and P. polare.
  • P. gallinacium is related to P. griffithsi
  • P. relictum is related to P. cathemerium, P. giovannolai and P. matutinum. P. relictum may be difficult to distinguish from P. giovannolai on either morphological grounds or on the basis of host species.
  • P. hexamerium is related to P. vaughni.
  • P. ashfordi is related to P. vaughni.

Vectors of avian malaria[edit]



Notes:

Sporogeny of P. circumflexum but not transmission has been recorded in Mansonia perturbans.

Avian malaria notes[edit]

  • P. relictum is known to infect over 70 bird families and 359 wild bird species so the record here should be regarded as incomplete. Additional host species can be found under the link Plasmodium relictum. It is likely that this species has been responsible for more bird extinctions than any other protist.
  • P. inconstans, P. irae, P. praecox, P. subpraecox and P. wasielewski have been re classified as P. relictum. P. subpraecox was described by Grassi and Feletti in 1892. P. wasielewski was described by Brumpt in 1909.
  • P. dominicana is species known only from fossil amber.[14] It is thought to have been a species infecting birds. It has been placed in the subgenus Nyssorhynchus.
  • The taxonomic status of P. corradettii (Laird, 1998) is currently regarded as dubious and may be revised.
  • There are currently 13 species recognised in the subgenus Novyella all of which are listed here.

A number of additional species have been described in birds - P. centropi, P. chloropsidis, P. gallinuae, P. herodialis, P. heroni, P. mornony, P. pericorcoti and P. ploceii - but the suggested speciation was based at least in part on the idea - 'one host - one species'. It has not been possible to reconcile the descriptions with any of the currently recognised species, and these are not currently regarded as valid species. As further investigations are made into this genus these species may be resurrected.

A species P. japonicum has been reported[15] but this appears to be the only report of this species and should therefore be regarded of dubious validity.

References[edit]

  1. ^ Wiersch S.C., Maier W.A., Kampen H. (2005) Plasmodium (Haemamoeba) cathemerium gene sequences for phylogenetic analysis of malaria parasites. Parasitol. Res. 96(2): 90-94
  2. ^ Martinsen E.S.,Waite J.L.,Schall J.J. Morphologically defined subgenera of Plasmodium from avian hosts: test of monophyly by phylogenetic analysis of two mitochondrial genes (2006) Parasitol. 1-8
  3. ^ Valkiūnas G., Zehtindjiev P., Hellgren O., Ilieva M., Iezhova T.A., Bensch S. (2007) Linkage between mitochondrial cytochrome b lineages and morphospecies of two avian malaria parasites, with a description of Plasmodium (Novyella) ashfordi sp. nov. Parasitol. Res.
  4. ^ Landau I, Chabaud AG, Bertani S, and Snounou G. (2003) Parassitologia. 45(3-4):119-123 Taxonomic status and re-description of Plasmodium relictum (Grassi et Feletti, 1891), Plasmodium maior Raffaele, 1931, and description of P. bigueti n. sp. in sparrows.
  5. ^ Kirkpatrick CE, Lauer DM. (1985) Hematozoa of raptors from southern New Jersey and adjacent areas. J Wildl. Dis. 21(1):1-6.
  6. ^ Earle RA, Horak IG, Huchzermeyer FW, Bennett GF, Braack LE, Penzhorn BL. (1991) The prevalence of blood parasites in helmeted guineafowls, Numida meleagris, in the Kruger National Park. Onderstepoort J. Vet. Res. 58(3):145-147.
  7. ^ Valkiūnas G., Zehtindjiev P., Dimitrov D., Krizanauskiene A., Iezhova T.A., Bensch S. (2008) Polymerase chain reaction-based identification of Plasmodium (Huffia) elongatum, with remarks on species identity of haemosporidian lineages deposited in GenBank. Parasitol. Res. 102(6):1185-1193.
  8. ^ a b Baillie SM, Brunton DH (2011) Diversity, distribution and biogeographical origins of Plasmodium parasites from the New Zealand bellbird (Anthornis melanura). Parasitology 9:1-9
  9. ^ Murata K., Nii R., Sasaki E., Ishikawa S., Sato Y., Sawabe K., Tsuda Y., Matsumoto R., Suda A., Ueda M. (2008) Plasmodium (Bennettinia) juxtanucleare infection in a captive white eared-pheasant (Crossoptilon crossoptilon) at a Japanese zoo. J. Vet. Med. Sci. 70(2):203-205
  10. ^ Christensen B.M., Barnes H.J., Rowley W.A. (1983) Vertebrate host specificity and experimental vectors of Plasmodium (Novyella) kempi sp. n. from the eastern wild turkey in Iowa. J. Wildl. Dis. 19(3):204-213
  11. ^ Manwell R.D. (1968) Plasmodium octamerium n. sp., an avian malaria parasite from the pintail whydah bird Vidua macroura. J. Protozool. 15(4):680-685
  12. ^ Valkiunas G., Iezhova T.A. (2001) A comparison of the blood parasites in three subspecies of the yellow wagtail Motacilla flava. J. Parasitol. 87(4):930-934.
  13. ^ Zehtindjiev P, Križanauskienė A, Bensch S, Palinauskas V, Asghar M, Dimitrov D, Scebba S, Valkiunas G (2012) A new morphologically distinct avian malaria parasite that fails detection by established PCR-based protocols for amplification of the cytochrome B gene. J Parasitol
  14. ^ Poinar G. (2005) Plasmodium dominicana n. sp. (Plasmodiidae: Haemospororida) from Tertiary Dominican amber. Systematic Parasitol. 61 (1) 47-52
  15. ^ Manwell R.D. (1966) Plasmodium japonicum, P. juxtanucleare and P. nucleophilum in the Far East. J. Protozool. 13(1):8-11.
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Giovannolaia

Giovanolaia is a subgenus of the genus Plasmodium created by Corradetti et al. in 1963.[1] The parasites within this subgenus infect birds.

This subgenus was shown on the basis of DNA analysis to be polyphyletic. A revision of this subgenus on a morphological basis by Landua et al. moved several of the species in this subgenus into a new subgenus Papernaia.[2]

Diagnostic features[edit]

Species in the subgenus Giovanolaia have the following characteristics:

Schizonts contain plentiful cytoplasm, are larger than the host cell nucleus and frequently displace it. They are found only in mature erythrocytes.

Gametocytes are elongated.

Both gametocytes and schizonts are stretched along the red cell nucleus.

Exoerythrocytic schizogony occurs in the mononuclear phagocyte system.

Species in this subgenus[edit]

References[edit]

  1. ^ Corradetti A., Garnham P.C.C., Laird M. (1963). "New classification of the avian malaria parasites". Parassitologia 5: 1–4. 
  2. ^ Landau I, Chavatte JM, Peters W, Chabaud A (March 2010). "The sub-genera of avian Plasmodium". Parasite 17 (1): 3–7. doi:10.1051/parasite/2010171003. PMID 20387732. 
  • "Plasmodium (Giovannolaia)". NCBI Taxonomy Browser. 418112. 


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Plasmodium

A plasmodium is also the macroscopic form of slime moulds.

Plasmodium, commonly known as the malaria parasite, is a large genus of parasitic protozoa. Infection with these protozoans is known as malaria, a deadly disease widespread in the tropics. The parasite always has two hosts in its life cycle: a mosquito vector and a vertebrate host.

The life-cycle is very complex, involving a sequence of different stages both in the vector and the host. These stages include sporozoites which are injected by the mosquito vector into the host's blood; latent hypnozoites which may rest undetected in the liver for up to 30 years; merosomes and merozoites which infect the red cells (erythrocytes) of the blood; trophozoites which grow in the red cells, and schizonts which divide there, producing more merozoites which leave to infect more red cells; and male and female sexual forms, gametocytes, which are taken up by other mosquitoes. In the mosquito's midgut, the gametocytes develop into gametes which fertilize each other to form motile zygotes which escape the gut, only to grow into new sporozoites which move to the mosquito's salivary glands, from where they are injected into the mosquito's next host, infecting it and restarting the cycle.

The genus Plasmodium was first described in 1885. It now contains about 200 species divided into several subgenera; as of 2006 the taxonomy was shifting, and species from other genera are likely to be added to Plasmodium. At least ten species infect humans; other species infect other animals, including birds, reptiles and rodents, while 29 species infect non-human primates. The parasite is thought to have originated from Dinoflagellates, photosynthetic protozoa.

The most common forms of human malaria are caused by Plasmodium falciparum, Plasmodium vivax, Plasmodium ovale, Plasmodium knowlesi, and Plasmodium malariae. P. falciparum malaria, common in sub-Saharan Africa, is especially dangerous.

Taxonomy and host range[edit]

The genus Plasmodium was created in 1885 by Marchiafava and Celli and there are over 200 species recognized. New species continue to be described.[1]

As of 2006, the genus is in need of reorganization as it has been shown that parasites belonging to the genera Haemocystis and Hepatocystis appear to be closely related to Plasmodium. It is likely that other species such as Haemoproteus meleagridis will be included in this genus once it is revised.

Host range among the mammalian orders is non uniform. At least 29 species infect non-human primates; rodents outside the tropical parts of Africa are rarely affected; a few species are known to infect bats, porcupines and squirrels; carnivores, insectivores and marsupials are not known to act as hosts.

In 1898 Ronald Ross demonstrated the existence of Plasmodium in the wall of the midgut and salivary glands of a Culex mosquito. For this discovery he won the Nobel Prize in 1902. However credit must also be given to the Italian professor Giovanni Battista Grassi, who showed that human malaria could only be transmitted by Anopheles mosquitoes. For some species the vector may not be a mosquito.

Mosquitoes of the genera Culex, Anopheles, Culiseta, Mansonia and Aedes may act as vectors. The known vectors for human malaria (more than 100 species) belong to the genus Anopheles. Bird malaria is commonly carried by species belonging to the genus Culex. Only female mosquitoes bite. Aside from blood both sexes live on nectar, but one or more blood meals are needed by the female for egg laying, because there is very little protein in nectar.

Life cycle[edit]

The life cycle of malaria parasites. A mosquito causes an infection by a bite. First, sporozoites enter the bloodstream, and migrate to the liver. They infect liver cells, where they multiply into merozoites, rupture the liver cells, and return to the bloodstream. Then, the merozoites infect red blood cells, where they develop into ring forms, trophozoites and schizonts that in turn produce further merozoites. Sexual forms are also produced, which, if taken up by a mosquito, will infect the insect and continue the life cycle.

The life cycle of Plasmodium is very complex. Sporozoites from the saliva of a biting female mosquito are transmitted to either the blood or the lymphatic system[2] of the recipient. The sporozoites then migrate to the liver and invade hepatocytes. This latent or dormant stage of the Plasmodium sporozoite in the liver is called the hypnozoite.

The development from the hepatic stages to the erythrocytic stages has been obscure. In 2006[3] it was shown that the parasite buds off the hepatocytes in merosomes containing hundreds or thousands of merozoites. These merosomes have been subsequently shown[4] to lodge in the pulmonary capillaries and to disintegrate there slowly over 48–72 hours releasing merozoites. Erythrocyte invasion is enhanced when blood flow is slow and the cells are tightly packed: both of these conditions are found in the alveolar capillaries.

Within the erythrocytes the merozoite grow first to a ring-shaped form and then to a larger trophozoite form. In the schizont stage, the parasite divides several times to produce new merozoites, which leave the red blood cells and travel within the bloodstream to invade new red blood cells. Most merozoites continue this replicative cycle, but some merozoites differentiate into male or female sexual forms (gametocytes) (also in the blood), which are taken up by the female mosquito.

In the mosquito's midgut, the gametocytes develop into gametes and fertilize each other, forming motile zygotes called ookinetes. The ookinetes penetrate and escape the midgut, then embed themselves onto the exterior of the gut membrane. Here they divide many times to produce large numbers of tiny elongated sporozoites. These sporozoites migrate to the salivary glands of the mosquito where they are injected into the blood of the next host the mosquito bites. The sporozoites move to the liver where they repeat the cycle.

The pattern of alternation of sexual and asexual reproduction is common in parasitic species. The evolutionary advantages of this type of life cycle were recognised by Mendel. Under favourable conditions, asexual reproduction is superior to sexual as the parent is well adapted to its environment and its descendents share all its genes. Transferring to a new host or in times of stress, sexual reproduction is generally superior as it shuffles the genes of two parents, producing a variety of individuals, some of which will be better adapted to the new environment.

Reactivation of the hypnozoites has been reported for up to 30 years after the initial infection in humans. The factors precipating this reactivation are not known. In the species Plasmodium malariae, Plasmodium ovale and Plasmodium vivax hypnozoites have been shown to occur. Reactivation does not occur in infections with Plasmodium falciparum. It is not known if hypnozoite reactivaction may occur with any of the remaining species that infect humans but this is presumed to be the case.

Evolution[edit]

The life cycle of Plasmodium is best understood in terms of its evolution.

The Apicomplexa—the phylum to which Plasmodium belongs—are thought to have originated within the Dinoflagellates, a large group of photosynthetic protozoa. It is thought that the ancestors of the Apicomplexa were originally prey organisms that evolved the ability to invade the intestinal cells and subsequently lost their photosynthetic ability. Some extant dinoflagelates, however, can invade the bodies of jellyfish and continue to photosynthesize, which is possible because jellyfish bodies are almost transparent. In other organisms with opaque bodies this ability would most likely rapidly be lost.

It is thought that Plasmodium evolved from a parasite spread by the orofaecal route which infected the intestinal wall. At some point this parasite evolved the ability to infect the liver. This pattern is seen in the genus Cryptosporidium, to which Plasmodium is distantly related. At some later point this ancestor developed the ability to infect blood cells and to survive and infect mosquitoes. Once mosquito transmission was firmly established the previous orofecal route of transmission was lost.

Current (2007) theory suggests that the genera Plasmodium, Hepatocystis and Haemoproteus evolved from Leukocytozoon species. Parasites of the genus Leukocytozoan infect white blood cells (leukocytes), liver and spleen cells and are transmitted by 'black flies' (Simulium species) — a large genus of flies related to the mosquitoes.

Leukocytes, hepatocytes and most spleen cells actively phagocytose particulate matter, making entry into the cell easier for the parasite. The mechanism of entry of Plasmodium species into erythrocytes is still very unclear, taking as it does less than 30 seconds. It is not yet known if this mechanism evolved before mosquitoes became the main vectors for transmission of Plasmodium.

Plasmodium evolved about 130 million years ago. This period coincided with the rapid spread of the angiosperms (flowering plants). This expansion in the angiosperms is thought to be due to at least one genomic duplication event. It seems probable that the increase in the number of flowers led to an increase in the number of mosquitoes and their contact with vertebrates.

Mosquitoes evolved in what is now South America about 230 million years ago. There are over 3500 species recognised but to date their evolution has not been well worked out so a number of gaps in our knowledge of the evolution of Plasmodium remain. It seems probable that birds were the first group infected by Plasmodium followed by the reptiles — probably the lizards. At some point primates and rodents became infected. The remaining species infected outside these groups seem likely to be due to relatively recent events.

P. falciparum, the most lethal malaria parasite of humans, evolved from a "nearly identical" parasite of western gorillas, not from chimpanzees, bonobos or ancient human populations.[5]

Molecular biology[edit]

Plasmodium is a Eukaryote, an organism whose cells have a nucleus, but with unusual features

All the species examined to date have 14 chromosomes, one mitochondrion and one plastid (an organelle similar to a chloroplast). The chromosomes vary from 500 kilobases to 3.5 megabases in length. It is presumed that this is the pattern throughout the genus. The plastid, unlike those found in algae, is not photosynthetic. Its function is not known but there is some suggestive evidence that it may be involved in reproduction.

On a molecular level, the parasite damages red blood cells using plasmepsin enzymes — aspartic acid proteases which degrade hemoglobin.

Taxonomy[edit]

Plasmodium belongs to the family Plasmodiidae (Levine, 1988), order Haemosporidia and phylum Apicomplexa. There are 450 recognised species in this order. Many species of this order are undergoing reexamination of their taxonomy with DNA analysis. It seems likely that many of these species will be reassigned after these studies have been completed.[6][7] For this reason the entire order is outlined here.

The genera Plasmodium, Fallisia and Saurocytozoon all cause malaria in lizards. All are carried by Diptera (true two-winged flies). Pigment is absent in the Garnia. Non pigmented gametocytes are typically the only forms found in Saurocytozoon: pigmented forms may be found in the leukocytes occasionally. Fallisia produce non pigmented asexual and gametocyte forms in leukocytes and thrombocytes.

Subgenera[edit]

The full taxonomic name of a species includes the subgenus but this is often omitted. The full name indicates some features of the morphology and type of host species.

The only two species in the sub genus Laverania are P. falciparum and P. reichenowi.

Species infecting monkeys and apes (the higher primates) with the exceptions of P. falciparum and P. reichenowi are classified in the subgenus Plasmodium.

Parasites infecting other mammals including lower primates (lemurs and others) are classified in the subgenus Vinckeia. The distinction between P. falciparum and P. reichenowi and the other species infecting higher primates was based on morphological findings but have since been confirmed by DNA analysis. Vinckeia, while previously considered to be something of a taxonomic 'rag bag', has been recently shown to form a coherent grouping. The remaining groupings here are based on the morphology of the parasites. Revisions to this system are likely as more species are subject to DNA analysis.

The four subgenera Giovannolaia, Haemamoeba, Huffia and Novyella were created by Corradetti et al.[8] for the known avian malarial species. A fifth — Bennettinia — was created in 1997 by Valkiunas.[9] The relationships between the subgenera are a matter of current investigation. Martinsen et al. 's recent (2006) paper outlines what was known at the time.[10]

As of 2007, P. juxtanucleare is the only known member of the subgenus Bennettinia.

Unlike the mammalian and bird malarias those affecting reptiles have been more difficult to classify. In 1966 Garnham classified those with large schizonts as Sauramoeba, those with small schizonts as Carinamoeba and the single then known species infecting snakes (Plasmodium wenyoni) as Ophidiella.[11] He was aware of the arbitrariness of this system and that it might not prove to be biologically valid. Telford in 1988 used this scheme as the basis for the accepted (2007) system.[12]

Species infecting humans[edit]

Trophozoites of the Plasmodium vivax parasite among human red blood cells

The species of Plasmodium that infect humans include:

The first four listed here are the most common species that infect humans. Nearly all human deaths from malaria are caused by the first species, P. falciparum, mainly in sub-Saharan Africa. With the use of the polymerase chain reaction additional species have been and are still being identified that infect humans.

One possible experimental infection has been reported with Plasmodium eylesi. Fever and low grade parasitemia were apparent at 15 days. The volunteer (Dr Bennett) had previously been infected by Plasmodium cynomolgi and the infection was not transferable to a gibbon (P. eylesi 's natural host) so this cannot be regarded as definitive evidence of its ability to infect humans. A second case has been reported that may have been a case of P. eylesi but the author was not certain of the infecting species.[13]

A possible infection with Plasmodium tenue has been reported.[14] This report described a case of malaria in a three-year-old black girl from Georgia, US, who had never been outside the US. She suffered from both P. falciparum and P. vivax malaria and while forms similar to those described for P. tenue were found in her blood even the author was skeptical about the validity of the diagnosis.

Confusingly, P. tenue was proposed in the same year (1914) for a species found in birds. The human species is now considered probably a misdiagnosis, and the bird species is described on the P. tenue page.

The only known host of P. falciparum and P, malariae is humans. P. vivax however can infect chimpanzees. Infection tends to be low grade but may be persistent and remain as source of parasites for humans for some time. P. vivax can also infect orangutans.[15]

P. ovale can be transmitted to chimpanzees. P. ovale has an unusual distribution, being found in Africa, the Philippines and New Guinea. In spite of its admittedly poor transmission to chimpanzees given its discontigous spread, it is suspected that P. ovale is a zoonosis with an as yet unidentified host. If so, the host is likely to be a primate. The remaining species capable of infecting humans all have other primate hosts.

Plasmodium shortii and Plasmodium osmaniae are now considered junior synonyms of Plasmodium inui

Taxonomy in parasitology before DNA based methods was always problematic, and revisions are continuing, leaving many obsolete names for Plasmodium species that infect humans.[16]

Infections in primates[edit]

Species of Plasmodium infect many primates across the world, such as the brown lemur, Eulemur fulvus, of Madagascar.

The species that infect primates other than humans include: P. bouillize, P. brasilianum, P. bucki, P. cercopitheci,P. coatneyi, P. coulangesi, P. cynomolgi, P. eylesi, P. fieldi, P. foleyi, P. fragile, P. girardi, P. georgesi, P. gonderi, P. hylobati, P. inui, P. jefferyi, P. joyeuxi, P. knowlesi, P. lemuris, P. percygarnhami, P. petersi, P. reichenowi, P. rodhaini, P. sandoshami, P. semnopitheci, P. silvaticum, P. simiovale, P. simium, P. uilenbergi, P. vivax and P. youngei.

Most if not all Plasmodium species infect more than one host: the host records shown here should be regarded as incomplete.

Infections in non-primate mammals[edit]

Many non-primate mammals, such as mouse-deer (Tragulus kanchil) can carry malaria parasites.

The subgenus Vinckeia was created by Garnham to accommodate the mammalian parasites other than those infecting primates. Species infecting lemurs have also been included in this subgenus.

P. aegyptensis, P. bergei, P. chabaudi, P. inopinatum, P. yoelli and P. vinckei infect rodents. P. bergei, P. chabaudi, P. yoelli and P. vinckei have been used to study malarial infections in the laboratory. Other members of this subgenus infect other mammalian hosts.

Infections in birds[edit]

Many bird species, from raptors to passerines like the red-whiskered bulbul (Pycnonotus jocosus), can carry malaria.

Species in five Plasmodium subgenera infect birdsBennettinia, Giovannolaia, Haemamoeba, Huffia and Novyella.[27] Giovannolaia appears to be a polyphyletic group and may be sudivided in the future.[10] DNA evidence is in 2014 helping to improve understanding of the diversity of Plasmodium species that infect birds.[28]

Infections in reptiles[edit]

Over 3000 species of lizard, including the Carolina anole (Anolis carolinensis), carry some 90 kinds of malaria.

Species in the subgenera Asiamoeba, Carinamoeba, Lacertaemoba, Paraplasmodium and Sauramoeba infect reptiles.[40]

Over 90 species and subspecies of Plasmodium infect lizards and they have been reported from over 3200 species of lizard and 29 species of snake. Only three species — P. pessoai, P. tomodoni and P. wenyoni — infect snakes.

Species reclassified into other genera[edit]

Further information: Hepatocystis and Garnia (protist)

References[edit]

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  42. ^ Garnham PC, Telford SR (November 1984). "A new malaria parasite Plasmodium (Sauramoeba) heischi in skinks (Mabuya striata) from Nairobi, with a brief discussion of the distribution of malaria parasites in the family Scincidae". J. Protozool. 31 (4): 518–21. doi:10.1111/j.1550-7408.1984.tb05494.x. PMID 6512723. 
  43. ^ Telford SR (October 1986). "Fallisia parasites (Haemosporidia: Plasmodiidae) from the flying lizard, Draco maculatus (Agamidae) in Thailand". J. Parasitol. 72 (5): 766–9. doi:10.2307/3281471. PMID 3100759. 
  44. ^ a b Telford SR (1979). "A taxonomic revision of small neotropical saurian Malarias allied to Plasmodium minasense". Ann Parasitol Hum Comp 54 (4): 409–22. PMID 533109. 
  45. ^ Telford SR, Telford SR (April 2003). [0362:RAROPP2.0.CO;2 "Rediscovery and redescription of Plasmodium pifanoi and description of two additional Plasmodium parasites of Venezuelan lizards"]. J. Parasitol. 89 (2): 362–8. doi:10.1645/0022-3395(2003)089[0362:RAROPP]2.0.CO;2. PMID 12760655. 

Further reading[edit]

Identification[edit]

  • Garnham, P.C.C. (1966). Malaria Parasites And Other Haemosporidia. Oxford: Blackwell. ISBN 0397601328. 
  • Hewitt, R.I. (1940). Bird Malaria. American Journal of Hygiene 15. Baltimore: Johns Hopkins Press. 
  • Laird, M. (1998). Avian Malaria in the Asian Tropical Subregion. Singapore: Springer. ISBN 9813083190. 

Biology[edit]

History[edit]

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Lacertaemoba

Lacertamoeba is a subgenus of the genus Plasmodium - all of which are parasitic protozoa. All species in this subgenus infect reptiles.

This subgenus was created by Telford to refine the classification of species then given as Plasmodium tropiduri. [1]

Diagnostic features[edit]

Species in the subgenus Lacertamoeba have the following characteristics:

The gametocytes are medium-sized

The schizonts undergo 3 to 5 nuclear divisions and are also medium-sized.

Species in this subgenus[edit]

References[edit]

  1. ^ Telford S. (1979) A taxonomic reconsideration of some Plasmodium species from iguanid lizards. Annales de parasitologie Humanie et Comparée 54: 129-144



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Huffia

Huffia is a subgenus of the genus Plasmodium - all of which are parasitic protozoa. The subgenus was created in 1963 by Corradetti et al..[1]

Species in this subgenus infect birds with malaria.

Diagnostic features[edit]

Species in the subgenus Huffia have the following characteristics:

  • Mature schizonts, while varying in shape and size, contain plentiful cytoplasm and are commonly found in immature erthryocytes.
  • Gametocytes are elongated.

Species in this subgenus[edit]

References[edit]

  1. ^ Corradetti A., Garnham P. C. C. and Laird M. (1963) New classification of the avian malaria parasites. Parassitologia 5, 1–4
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Plasmodium species infecting primates


Red blood cell infected with malaria

The Plasmodium species infecting primates include the parasites causing malaria in humans.

Species infecting humans[edit]

(Plasmodium brasilianum and Plasmodium rhodiani which have been reported to infect humans, are likely synonymous with P. malariae)

The first six listed here are the only common species that infect humans. While infection of humans by other species is known, they are quite rare, in some instances, only a single case. In a number of the cases, the means of infection is unknown, and may be due to accident, i.e. infection by laboratory equipment or a bite by an animal.

With the use of the polymerase chain reaction additional species have been and are still being identified that infect humans:

One possible experimental infection has been reported with Plasmodium eylesi. Fever and low grade parasitemia were apparent at 15 days. The volunteer (Dr Bennett) had previously been infected by Plasmodium cynomolgi and the infection was not transferable to a gibbon (P. eylesi 's natural host) so this cannot be regarded as definitive evidence of its ability to infect humans. A second case has been reported that may have been a case of P. eylesi but the author was not certain of the infecting species.[4]

A possible infection with Plasmodium tenue has been reported.[5] This report described a case of malaria in a three year old black girl from Georgia, USA who had never been outside the US. She suffered from both P. falciparum and P. vivax malaria and while forms similar to those described for P. tenue were found in her blood even the author was skeptical about the validity of the diagnosis.

Confusingly Plasmodium tenue was proposed in the same year (1914) for a species found in birds. The human species is now considered to be likely to have been a misdiagnosis and the bird species is described on the Plasmodium tenue page.

Notes[edit]

falciparum

Until recently the only known host of P. falciparum was humans but this species has also been described in gorillas (Gorilla gorilla)[6] and bonbos[7] There has been a single report of P. falciparum in a brown howler monkey (Alouatta guariba) and in black howler monkeys (Alouatta caraya)[8] but until this is confirmed its validity should be considered dubious.

A possible report of P. falciparum in a greater spot-nosed monkeys (Cercopithecus nictitans) has not been confirmed in a large survey.[9]

A species that clusters with P. falciparum and P. reichenowi has been identified in Gabon, Africa in chimpanzees (Pan troglodytes).[10] This appears to have diverged from these two species about 21 million years ago. It has only been identified from the sequence of its mitochondrion to date and further work is needed to characterise the species. A second report has confirmed the existence of this species in chimpanzees.[6] A third report has confirmed the existence of this species.[11]

Night monkeys (Aotus nigriceps) can be infected with P. falciparum. This infection may occur naturally.[12] Their potential role - if any - as a source of human infection is unknown.

Two additional species within the subgenus Laverania have been identified on the basis of DNA sequences alone: Plasmodium billbrayi and Plasmodium billcollinsi.[6] and bonbos[7] P. billbrayi was found in two subspecies of chimpanzee (Pan troglodytes troglodytes and Pan troglodytes schweinfurthii). P. billcollinsi was found in only one subspecies of chimpanzee (Pan troglodytes troglodytes). Further work is needed to characterise these species.

malariae

Humans are currently considered to be the only host for P. malariae. However Rodhain and Dellaert in the 1940s showed with transmission studies that P. malariae was present in chimpanzees.[13][14] The presence of P. malaria in chimpanzees has been reported in Japan suggesting that this species may be able to act as a host.[15] A second paper has described the presence of P. malaria in wild chimpanzees.[11] Another paper has reported several cases of P. malariae in brown howler monkey (Alouatta guariba) and black howler monkeys (Alouatta caraya)[8] It has been shown that splectomised three-striped night monkey (Aotus trivirgatus) can be infected with P. malariae.[16] Another paper has confirmed the presence of P. malaria in chimpanzees.[17]

The existence of multiple independent reports seem to suggest that the chimpanzee and possibly other species may act as a host to P. malaria at least occasionally.

vivax

P. vivax will infect chimpanzees. Infection tends to be low grade but may be persistent and remain as source of parasites for humans for some time. P. vivax is also known to infect orangutans[18] and the brown howler monkey (Alouatta guariba clamitans)[8] P. vivax has been reported from chimpanzees living in the wild.[11] It has been suggested that vivax infection of the great apes in Africa may act as a reservoir given the prevalence of Duffy antigen negative humans in this area.[19]

ovale

Like P. vivax, P. ovale has been shown to be transmittable to chimpanzees. P. ovale has an unusual distribution pattern being found in Africa, Myanmar the Philippines and New Guinea. In spite of its admittedly poor transmission to chimpanzees given its discontigous spread, it is suspected that P. ovale may in fact be a zooenosis with an as yet unidentified host. If this is actually the case, the host seems likely to be a primate. A report has been published suggesting that P. ovale may be a natural parasite of chimpanzees[20] but this needs confirmation. P. ovale has since been described from chimpanzees living in the wild.[11] This suggests that human infection with this species may as previously suspected be a zoonosis.

It has been recently shown that P. ovale is actually two genetically distinct species that coexist. These species are Plasmodium ovale curtisi and Plasmodium ovale wallikeri.[21] These two species separated between 1.0 and 3.5 million years ago.

knowlesi

Plasmodium knowlesi has a natural reservoir in the macaques of Southeast Asia, and was only in 1965 identified as being transmissible to humans.

Other species

The remaining species capable of infecting humans all have other primate hosts.

Plasmodium taxonomy[edit]

  • P. cynomolgi - P. cynomolgi bastianelli, P. cynomolgi ceylonensis and P. cynomolgi cynomolgi.
  • P. inui - P. inui inui and P. inui shortii
  • P. knowlesi - P. knowlesi edesoni and P. knowlesi knowlesi.
  • P. ovale - P. ovale curtisi and P. ovale wallikeri
  • P. vivax - P. vivax hibernans, P. vivax chesson and P. vivax multinucleatum.

Interrelatedness - The evolution of these species is still being worked out and the relationships given here should be regarded as tentative. This grouping, while originally made on morphological grounds, now has considerable support at the DNA level.

  • P. brasilianum, P. inui and P. rodhaini are similar to P. malariae (quartan malaria group)
  • P. cynomolgi, P. fragile, P. knowlesi, P. simium and P. schwetzi are similar to P. vivax
  • P. fieldi and P. simiovale are similar to P. ovale
  • P. falciparum is closely related to P. reichenowi.

Notes[edit]

  • P. kochi has been described as a parasite of monkeys. This species is currently classified as Hepatocystis kochi. This may be subject to revision.
  • P. brasilianum and P. rodhaini seem likely to be the same species as P. malariae.
  • P. lemuris may actually belong to the Haemoproteus genus. Clarification of this point awaits DNA examination.
  • P. shortii is currently (2007) regarded as a junior synonym of P. inui.

Species previously described as infecting humans but no longer recognised as valid[edit]

Taxonomy in parasitology until the advent of DNA based methods has always been a problem and revisions in this area are continuing. A number of synonyms have been given for the species infecting humans that are no longer recognised as valid.[22] Since perusal of the older literature may be confusing some currently defunct species names are listed here.

P. camerense

P. causiasium
P. golgi
P. immaculatum
P. laverani var. tertium
P. laverani var. quartum
P. malariae var. immaculatum
P. malariae var. incolor
P. malariae var. irregularis
P. malariae var. parva
P. malariae var. quartanae
P. malariae var. quotidianae
P. perniciosum
P. pleurodyniae
P. praecox
P. quartana
P. quotidianum
P. sedecimanae
P. tenue
P. undecimanae
P. vegesio-tertaniae

P. vivax-minuta

Plasmodium shortii and Plasmodium osmaniae are now considered to be junior synonyms of Plasmodium inui.

Species infecting other hosts[edit]

Most if not all Plasmodium species infect more than one host: the host records shown here should be regarded as incomplete.

  • P. malagasi - lemurs
  • P. reichenowi - chimpanzee (Pan) species[11] and gorilla (Gorilla) species
  • P. rodhaini - chimpanzee (Pan) species and gorilla (Gorilla) species
  • P. schwetzi - chimpanzee (Pan) species and gorilla (Gorilla) species

It has been proposed that the species P. gora and P. gorb should be renamed P. adleri and P. blacklocki respectively.

Primate groups and Plasmodium species[edit]

New World monkeys of the family Cebidae: P. brasilianum and P. simium

Old World monkeys of the Cercopithecidae family: P. coatneyi, P. cynomolgi, P. fieldi, P. fragile, P.gonderi, P. georgesi, P. inui, P. knowlesi, P. petersi, P. shortti and P. simiovale

Gibbons of the Hylobatidae family: P. eylesi, P. hylobati, P. jefferyi and P. youngi

Orangutans (Pongo): P. pitheci and P. silvaticum

Gorillas and chimpanzees: P. billcollini, P. billbrayii, P. falciparum, P. gabonensi, P. gora, P. gorb, P. reichenowi, P. rodhaini and P. schwetzi

Mosquitoes known to transmit human malaria listed by region[edit]

This listing may be incomplete as the taxonomy of this genus is under revision.

North American

Central American

South American

North Eurasian

Mediterranean

Afro-Arabian

Afrotropical

Indo-Iranian

Indo-Chinese hills

Malaysian

Chinese

Australasian

Primate mosquito vectors and associated Plasmodium species[edit]

References[edit]

  1. ^ Coatney GR, Chin W, Contacos PG, King HK (1966). "Plasmodium inui, a quartan-type malaria parasite of Old World monkeys transmissible to man". J Parasitol 52: 660–666. doi:10.2307/3276423. 
  2. ^ Contacos PG, Coatney GR, Orihel TC, Collins WE, Chin W (1970). "Transmission of Plasmodium schwetzi from the chimpanzee to man by mosquito bite". Am J Trop Med Hyg 19 (2): 190–5. PMID 5443069. 
  3. ^ Rodhain J, Dellaert R (1955). "Contribution a l'etude de Plasmodium schwetzi E. Brumpt (2eme note). Transmission de Plasmodium schwetzi a l'homme". Ann. Soc. Belg. Med. Trop. 35: 757–75. 
  4. ^ Tsukamoto M (1977). "An imported human malarial case characterized by severe multiple infections of the red blood cells". Ann. Trop. Med. Parasit. 19 (2): 95–104. 
  5. ^ Russel PF (1928). "Plasmodium tenue (Stephens): A review of the literature and a case report". Am. J. Trop. Med. s1–8 (5): 449–79. 
  6. ^ a b c Prugnolle F, Durand P, Neel C, Ollomo B, Ayala FJ, Arnathau C, Etienne L, Mpoudi-Ngole E, Nkoghe D et al. (2010). "African great apes are natural hosts of multiple related malaria species, including Plasmodium falciparum". Proc. Natl. Acad. Sci. USA 107 (4): 1458–1463. doi:10.1073/pnas.0914440107. PMC 2824423. PMID 20133889. 
  7. ^ a b Krief S, Escalante AA, Pacheco MA, Mugisha L, André C, Halbwax M, Fischer A, Krief JM, Kasenene JM, Crandfield M, Cornejo OE, Chavatte JM, Lin C, Letourneur F, Grüner AC, McCutchan TF, Rénia L, Snounou G (2010) On the Diversity of malaria parasites in African apes and the origin of Plasmodium falciparum from bonobos. PloS Pathog. 12;6(2):e1000765
  8. ^ a b c Duarte AM, Malafronte Rdos S, Cerutti C (August 2008). "Natural Plasmodium infections in Brazilian wild monkeys: Reservoirs for human infections?". Acta Trop. 107 (2): 179–85. doi:10.1016/j.actatropica.2008.05.020. PMID 18620330. 
  9. ^ Ayouba A, Mouacha F, Learn GH, Mpoudi-Ngole E, Rayner JC, Sharp PM, Hahn BH, Delaporte E, Peeters M (2012) Ubiquitous Hepatocystis infections, but no evidence of Plasmodium falciparum-like malarial parasites in wild greater spot-nosed monkeys (Cercopithecus nictitans). Int J Parasitol
  10. ^ Ollomo B, Durand P, Prugnolle F (May 2009). "A new malaria agent in African hominids". PLoS Pathog. 5 (5): e1000446. doi:10.1371/journal.ppat.1000446. PMC 2680981. PMID 19478877. 
  11. ^ a b c d e f g h i Kaiser M, Lowa A, Ulrich M, Ellerbrok H, Goffe AS, Blasse A, Zommers Z, Couacy-Hymann E, Babweteera F et al. (Dec 2010). "Wild chimpanzees infected with 5 Plasmodium species". Emerg Infect Dis 16 (12): 1956–1959. doi:10.3201/eid1612.100424. 
  12. ^ da Silva Araújo M, Messias MR, Figueiró MR, Gil LH, Probst CM, de Medeiros Vidal N, Katsuragawa TH, Krieger MA, Pereira da Silva LH, Ozaki LS (2013) Natural Plasmodium infection in monkeys in the state of Rondonia (Brazilian Western Amazon). Malar J 12(1):180
  13. ^ Rodhain J (1940) Les plasmodiums des anthropoids de l'Afrique centrale et leurs relations avec les plasmodiums humains. Récepticité de l'homme au Plasmodium malariae. (Plasmodium rodhaini Brumpt) du chimpanzé. C. R. Soc. Biol. 133:276-277
  14. ^ Rodhain J and Dellaert R (1943) L'infection á Plasmodium malariae du chimpanzé chez l'homme. Etude d'une premiére souche isolée de l'anthropoide Pan satyrus verus. Ann. Soc. Belge. Med. Trop. 23:19-46
  15. ^ Hayakawa T, Arisue N, Udono T (2009). "Identification of Plasmodium malariae, a human malaria parasite, in imported chimpanzees". PLoS ONE 4 (10): e7412. doi:10.1371/journal.pone.0007412. PMC 2756624. PMID 19823579. 
  16. ^ Collins WE, Contacos PG (1969). "Infectivity of Plasmodium malariae in the Aotus trivirgatus monkey to Anopheles freeborni mosquitoes". J Parasitol 55 (6): 1253–1257. doi:10.2307/3277270. 
  17. ^ Pacheco MA, Cranfield M, Cameron K, Escalante AA (2013) Malarial parasite diversity in chimpanzees: the value of comparative approaches to ascertain the evolution of Plasmodium falciparum antigens. Malar J 12(1):328
  18. ^ Reid MJ, Ursic R, Cooper D (December 2006). "Transmission of human and macaque Plasmodium spp. to ex-captive orangutans in Kalimantan, Indonesia". Emerging Infect. Dis. 12 (12): 1902–8. doi:10.3201/eid1212.060191. PMID 17326942. 
  19. ^ Prugnolle F, Rougeron V, Becquart P, Berry A, Makanga B, Rahola N, Arnathau C, Ngoubangoye B, Menard S, Willaume E, Ayala FJ, Fontenille D, Ollomo B, Durand P, Paupy C, Renaud F (2013) Diversity, host switching and evolution of Plasmodium vivax infecting African great apes. Proc Natl Acad Sci USA
  20. ^ Duval L, Nerrienet E, Rousset D (2009). "Chimpanzee malaria parasites related to Plasmodium ovale in Africa". PLoS ONE 4 (5): e5520. doi:10.1371/journal.pone.0005520. PMC 2677663. PMID 19436742. 
  21. ^ Sutherland CJ, Tanomsing N, Nolder D, Oguike M, Jennison C, Pukrittayakamee S, Dolecek C, Hien TT, do Rosário VE, Arez AP, Pinto J, Michon P, Escalante AA, Nosten F, Burke M, Lee R, Blaze M, Otto TD, Barnwell JW, Pain A, Williams J, White NJ, Day NP, Snounou G, Lockhart PJ, Chiodini PL, Imwong M, Polley SD (2010). "Two nonrecombining sympatric forms of the human malaria parasite Plasmodium ovale occur globally". J Infect Dis 201 (10): 1544–50. doi:10.1086/652240. PMID 20380562.