:''A 'plasmodium' is also the macroscopic form of the
protist known as a
slime mould.''
'''Plasmodium''' is a genus of parasitic
protozoa. Infection with this genus is known as
malaria. The parasite always has two hosts in its
life cycle: a
mosquito vector and a
vertebrate host. At least ten species infect humans. Other species infect other animals, including
birds,
reptiles and
rodents.
Taxonomy and host range
The
genus ''Plasmodium'' was created in
1885 by
Marchiafava and
Celli and there are over 175 species currently recognised. New species continue to be described.
[1]
The genus is currently (
2006) in need of reorganisation 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.
Life cycle
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. It is worth noting, however, that for some species the vector may not be a mosquito.
Mosquitoes of the genera ''
Culex'', ''
Anopheles'', ''
Culiceta'', ''
Mansonia'' and ''
Aedes'' may act as vectors. The currently known vectors for human malaria (> 100 species) all 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 as the protein content of nectar is very low. The life cycle of ''Plasmodium'' was discovered by Ross who worked with species from the genus ''Culex''.
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. From the hepatocytes, the parasite replicates into thousands of
merozoites, which then invade
red blood cells.
Here the parasite grows from a ring-shaped form to a larger
trophozoite form. In the 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.
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
This life cycle is best understood in terms of its
evolution. 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 is coincidental 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.
Presently 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.
At the present time (2007) DNA sequences are available from fewer than sixty species and most of these are from species infecting either rodent or primate hosts. The evolution proposed here should be regarded as speculative and subject to revision as data becomes available.
Reproduction
The pattern of alternation of sexual and asexual reproduction which may seem confusing at first is a very common pattern 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 these genes. Transferring to a new host or in times of stress, sexual reproduction is generally superior as this produces a shuffling of
genes which on average at a population level will produce individuals better adapted to the new environment.
Molecular biology
All the species examined to date have 14
chromosomes, one
mitochondrion and one
plastid. The chromsomes 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.
Diagnostic characteristics of the genus ''Plasmodium''
★ Forms gamonts in erythrocytes
★ Merogony occurs in erythrocytes and in other tissues
★ Hemozoin is present
★ Vectors are either mosquitos or sandflies
★ Vertebrate hosts include mammals, birds and reptiles
Taxonomy
''Plasmodium'' belongs to the
family Plasmodiidae (Levine,
1988),
order Haemosporidia and
phylum Apicomplexia. There are currently 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 re assigned after these studies have been completed.
[ Perkins SL, Schall JJ (2002) A molecular phylogeny of malarial parasites recovered from cytochrome b gene sequences. J. Parasitology 88 (5): 972-978 ][ Yotoko, K. S. C. and Elisei C. (2006) Malaria parasites (Apicomplexa, Haematozoea) and their relationships with their hosts: is there an evolutionary cost for the specialization? J. Zoo. Syst. Evol. Res. 44 (4) 265 ] For this reason the entire order is outlined here.
★ Order Haemosporida
★ Genus ''
Bioccala''
Family ''
Haemoproteidae''
★ Genus ''
Haemoproteus''
★
★ Subgenus ''
Parahaemoproteus''
★
★ Subgenus ''
Haemoproteus''
Family ''
Garniidae
★ Genus ''
Fallisia''
★
★ Subgenus ''
Plasmodioides''
Family ''
Leucocytozoidae''
★ Genus ''
Leukocytozoon''
★
★ Subgenus ''
Leucocytozoon''
★
★ Subgenus ''
Akiba''
Family ''
Plasmodiidae''
★ Genus ''
Billbraya''
★ Genus ''
Dionisia''
★ Genus ''
Hepatocystis''
★ Genus ''
Mesnilium''
★ Genus ''
Nycteria''
★ Genus ''Plasmodium''
★
★ Subgenus ''
Asiamoeba''
★
★ Subgenus ''
Bennettinia''
★
★ Subgenus ''
Carinamoeba''
★
★ Subgenus ''
Giovannolaia''
★
★ Subgenus ''
Haemamoeba''
★
★ Subgenus ''
Huffia''
★
★ Subgenus ''
Lacertaemoba''
★
★ Subgenus ''
Laverania''
★
★ Subgenus ''
Novyella''
★
★ Subgenus ''Plasmodium''
★
★ Subgenus ''
Paraplasmodium''
★
★ Subgenus ''
Sauramoeba''
★
★ Subgenus ''
Vinckeia''
★ Genus ''
Polychromophilus''
★ Genus ''
Rayella''
★ Genus ''
Saurocytozoon''
'Notes:'
The genera ''Plasmodium'', ''Fallisia'' and ''Saurocytozoon'' all cause malaria in lizards. All are carried by
Dipteria (roughly speaking the 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
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 the 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 - perhaps rather surprisingly - to form a coherent grouping.
The remaining groupings here are based on the morphology of the parasites. Revisions to this system are likely to occur in the future as more species are subject to analysis of their
DNA.
The four subgenera ''
Giovannolaia'', ''
Haemamoeba'', ''
Huffia'' and ''
Novyella'' were created by Corradetti et al
[3] for the known avian malarial species. A fifth - ''
Bennettinia'' - was created in 1997 by Valkiunas.
[4] The relationships between the subgenera are the matter of current investigation. Martinsen ''et al'' 's recent (
2006) paper outlines what is currently (
2007) known.
[5]
''P. juxtanucleare'' is currently (
2007) 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''.
[6] 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 currently accepted (2007) system.
[7]
'Classification criteria'
Species in the subgenus ''Bennettinia'' have the following characteristics:
★
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.
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.
★ Exoerythrocytic schizogony occurs in the
mononuclear phagocyte system.
Species in the subgenus ''Haemamoeba'' have the following characteristics:
★ 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.
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 the subgenus ''Novyella'' have the following characteristics:
★ Mature schisonts 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
Species in the subgenus ''
Carinamoeba'' have the following characteristics:
★ Infect lizards
★ Schizonts normally give rise to less than 8 merozoites
Species in the subgenus ''
Sauramoeba'' have the following characteristics:
★ Infect lizards
★ Schizonts normally give rise to more than 8 merozoites
'Notes'
★ The erythrocytes of both reptiles and birds retain their nucleus, unlike those of mammals. The reason for the loss of the nucleus in mammalian erythocytes remains unknown.
★ The presence of elongated gametocytes in several of the avian subgenera and in ''Laverania'' in addition to a number of clinical features suggested that these might be closely related. This is is no longer thought to be the case.
Species listed by subgenera
''Plasmodium (Asiamoeba) draconis''
''Plasmodium (Asiamoeba) vastator''
''Plasmodium (Bennettinia) juxtanucleare''
''Plasmodium (Carinamoeba) basilisci''
''Plasmodium (Carinamoeba) clelandi''
''Plasmodium (Carinamoeba) lygosomae''
''Plasmodium (Carinamoeba) mabuiae''
''Plasmodium (Carinamoeba) minasense''
''Plasmodium (Carinamoeba) rhadinurum''
''Plasmodium (Carinamoeba) volans''
''Plasmodium (Giovannolaia) anasum''
''Plasmodium (Giovannolaia) circumflexum''
''Plasmodium (Giovannolaia) dissanaikei''
''Plasmodium (Giovannolaia) durae''
''Plasmodium (Giovannolaia) fallax''
''Plasmodium (Giovannolaia) formosanum''
''Plasmodium (Giovannolaia) gabaldoni''
''Plasmodium (Giovannolaia) garnhami''
''Plasmodium (Giovannolaia) gundersi''
''Plasmodium (Giovannolaia) hegneri''
''Plasmodium (Giovannolaia) lophurae''
''Plasmodium (Giovannolaia) pedioecetii''
''Plasmodium (Giovannolaia) pinnotti''
''Plasmodium (Giovannolaia) polare''
''Plasmodium (Haemamoeba) cathemerium''
''Plasmodium (Haemamoeba) coggeshalli''
''Plasmodium (Haemamoeba) elongatum''
''Plasmodium (Haemamoeba) gallinaceum''
''Plasmodium (Haemamoeba) giovannolai''
''Plasmodium (Haemamoeba) lutzi''
''Plasmodium (Haemamoeba) matutinum''
''Plasmodium (Haemamoeba) paddae''
''Plasmodium (Haemamoeba) parvulum''
''Plasmodium (Haemamoeba) relictum''
''Plasmodium (Haemamoeba) tejera''
''Plasmodium (Huffia) elongatum''
''Plasmodium (Huffia) hermani''
''Plasmodium (Lacertaemoba) floridense''
''Plasmodium (Lacertaemoba) tropiduri''
''Plasmodium (Laverania) falciparum''
''Plasmodium (Laverania) reichenowi''
''Plasmodium (Novyella) ashfordi''
''Plasmodium (Novyella) bertii''
''Plasmodium (Novyella) bambusicolai''
''Plasmodium (Novyella) columbae''
''Plasmodium (Novyella) corradettii''
''Plasmodium (Novyella) dissanaikei''
''Plasmodium (Novyella) hexamerium''
''Plasmodium (Novyella) jiangi''
''Plasmodium (Novyella) kempi''
''Plasmodium (Novyella) nucleophilum''
''Plasmodium (Novyella) papernai''
''Plasmodium (Novyella) paranucleophilum''
''Plasmodium (Novyella) rouxi''
''Plasmodium (Novyella) vaughani''
''Plasmodium (Paraplasmodium) chiricahuae''
''Plasmodium (Paraplasmodium) mexicanum''
''Plasmodium (Paraplasmodium) pifanoi''
''Plasmodium (Plasmodium) bouillize''
''Plasmodium (Plasmodium) brasilianum''
''Plasmodium (Plasmodium) cercopitheci''
''Plasmodium (Plasmodium) coatneyi''
''Plasmodium (Plasmodium) cynomolgi''
''Plasmodium (Plasmodium) eylesi''
''Plasmodium (Plasmodium) fieldi''
''Plasmodium (Plasmodium) fragile''
''Plasmodium (Plasmodium) georgesi''
''Plasmodium (Plasmodium) girardi''
''Plasmodium (Plasmodium) gonderi''
''Plasmodium (Plasmodium) inui''
''Plasmodium (Plasmodium) jefferyi''
''Plasmodium (Plasmodium) joyeuxi''
''Plasmodium (Plasmodium) knowlei''
''Plasmodium (Plasmodium) hyobati''
''Plasmodium (Plasmodium) malariae''
''Plasmodium (Plasmodium) ovale''
''Plasmodium (Plasmodium) petersi''
''Plasmodium (Plasmodium) pitheci''
''Plasmodium (Plasmodium) rhodiani''
''Plasmodium (Plasmodium) schweitzi''
''Plasmodium (Plasmodium) semiovale''
''Plasmodium (Plasmodium) semnopitheci''
''Plasmodium (Plasmodium) silvaticum''
''Plasmodium (Plasmodium) simium''
''Plasmodium (Plasmodium) vivax''
''Plasmodium (Plasmodium) youngi''
''Plasmodium (Sauramoeba) achiotense''
''Plasmodium (Sauramoeba) adunyinkai''
''Plasmodium (Sauramoeba) aeuminatum''
''Plasmodium (Sauramoeba) agamae''
''Plasmodium (Sauramoeba) beltrani''
''Plasmodium (Sauramoeba) brumpti''
''Plasmodium (Sauramoeba) cnemidophori''
''Plasmodium (Sauramoeba) diploglossi''
''Plasmodium (Sauramoeba) giganteum''
''Plasmodium (Sauramoeba) heischi''
''Plasmodium (Sauramoeba) josephinae''
''Plasmodium (Sauramoeba) pelaezi''
''Plasmodium (Sauramoeba) zonuriae''
''Plasmodium (Vinckeia) achromaticum''
''Plasmodium (Vinckeia) aegyptensis''
''Plasmodium (Vinckeia) anomaluri''
''Plasmodium (Vinckeia) atheruri''
''Plasmodium (Vinckeia) berghei''
''Plasmodium (Vinckeia) booliati''
''Plasmodium (Vinckeia) brodeni''
''Plasmodium (Vinckeia) bubalis''
''Plasmodium (Vinckeia) bucki''
''Plasmodium (Vinckeia) caprae''
''Plasmodium (Vinckeia) cephalophi''
''Plasmodium (Vinckeia) chabaudi''
''Plasmodium (Vinckeia) coulangesi''
''Plasmodium (Vinckeia) cyclopsi''
''Plasmodium (Vinckeia) foleyi''
''Plasmodium (Vinckeia) girardi''
''Plasmodium (Vinckeia) inopinatum''
''Plasmodium (Vinckeia) lemuris''
''Plasmodium (Vinckeia) melanipherum''
''Plasmodium (Vinckeia) odocoilei''
''Plasmodium (Vinckeia) percygarnhami''
''Plasmodium (Vinckeia) sandoshami''
''Plasmodium (Vinckeia) traguli''
''Plasmodium (Vinckeia) tyrio''
''Plasmodium (Vinckeia) uilenbergi''
''Plasmodium (Vinckeia) vinckei''
''Plasmodium (Vinckeia) watteni''
''Plasmodium (Vinckeia) yoelli''
'Notes'
''
Ophidiella'' was a subgenus created by Garnham in
1966 for the species infecting snakes. Presently (2007) it is no longer in use.
Species infecting humans
Red blood cell infected with
malaria
The species of ''Plasmodium'' that parasitise humans include:
★ ''
Plasmodium falciparum'' (the cause of malignant tertian malaria)
★ ''
Plasmodium vivax'' (the most frequent cause of benign tertian malaria)
★ ''
Plasmodium ovale'' (the other, less frequent, cause of benign tertian malaria)
★ ''
Plasmodium malariae'' (the cause of benign quartan malaria)
★ ''
Plasmodium knowlesi''
★ ''
Plasmodium brasilianum''
★ ''
Plasmodium cynomolgi''
★ ''
Plasmodium cynomolgi bastianellii''
★ ''
Plasmodium inui''
★ ''
Plasmodium rhodiani''
★ ''
Plasmodium schweitzi''
★ ''
Plasmodium semiovale''
★ ''
Plasmodium simium''
The first four listed here are the most common species that infect humans. 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.
[8]
A possible infection with ''
Plasmodium tenue'' has been reported.
[9] 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':
The only known host of ''P. falciparum'' are humans; neither is any other host currently known for ''P. malariae.''
''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.
[10]
Like ''P. vivax'', ''P. ovale'' has been shown to be transmittable to
chimpanzees. ''P. ovale'' has a unusual distribution pattern 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'' 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.
The remaining species capable of infecting humans all have other primate hosts.
''Plasmodium shortii'' and ''Plasmodium osmaniae'' are now considered to be junior synonyms of ''Plasmodium inui''
'Species no longer recognised as valid'
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 synonoms have been given for the species infecting humans that are no longer recognised as valid.
[11] Since perusal of the older literature may be confusing some of these 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''
Infections in primates
'Listing'
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''.
'Host records' - Most if not all ''Plasmodium'' species infect more than one host: the host records shown here should be regarded as being incomplete.
★ ''P. bouillize'' - ''
Cercopithecis campbelli''
★ ''P. brasilianum'' - ''
Alouatta fusca'', ''
Alouatta palliata'', ''
Alouatta seniculus straminea'', ''
Alouatta villosa'', several
night monkey (''
Aotus'') species, ''
Ateles fusciceps'', ''
Ateles geoffroyi'', ''
Ateles geoffroyi grisescens'', ''
Ateles paniscus'', ''
Ateles paniscus paniscus'', ''
Ateles paniscus chamek'', ''
Brachyteles arachnoides'', ''
Callicebus moloch ornatus'', ''
Callicebus torquatus'', ''
Cebus albifrons'', ''
Cebus apella'', ''
Cebus capucinus'', ''
Cebus capucinus capucinus'', ''
Cebus capucinus imitator'', ''
Chiropotes chiropotes'', ''
Lagothrix cana'', ''
Lagothrix infumata'', ''
Lagothrix lagotricha'', ''
Saimiri boliviense'' and ''
Saimiri sciureus''.
★ ''P. bucki'' - ''
Lemur macaco macaco''
★ ''P. cercopitheci'' - ''
Cercopithecis nictitans''
★ ''P. coatneyi'' - several
macaque species: ''
Macaca fascicularis'' and ''
Macaca irus''.
★ ''P. coulangesi'' - ''
Lemur macaco macaco''
★ ''P. cynomolgi'' - ''
Macaca arctoides'', ''
Macaca cyclopis'', ''
Macaca fascicularis'', ''
Macaca mulatta'', ''
Macaca nemestrina'', ''
Macaca radiata'', ''
Macaca sinica'', ''
orangutans'' (''
Pongo''), ''
Presbytis cristatus'' and ''
Presbytis entellus''
★ ''P. eylesi'' - several
gibbon (''
Hylobates'') species including ''
Hylobates lar''
★ ''P. fieldi'' - ''
Macaca fascicularis'' and ''
Macaca nemestrina''
★ ''P. foleyi'' - ''
Lemur fulvus rufus''
★ ''P. fragile'' - several
macaque species - ''
Macaca fascicularis'', ''
Macaca mulatta'', ''
Macaca radiata'', and ''
Macaca sinica''
★ ''P. georgesi'' - ''
Cercocebus albigena''
★ ''P. girardi'' - ''
Lemur fulvus rufus'', ''
Lemur macaco macaco''
★ ''P. gonderi'' -
Mangabeys: ''
Cercocebus albigena'', ''
Cercocebus aterrimus'', ''
Cercocebus atys'', ''
Cercocebus galeritus agilus'' and
drills (''
Mandrillus leucophaeus'')
★ ''P. hylobati'' - several
gibbon (''
Hylobates'') species including ''
Hylobates lar'' and ''
Hylobates moloch''
★ ''P. inui'' - the
Celebes black
ape (''
Cynopithecus niger''), ''
Macaca fascicularis'', ''
Macaca mulatta'', ''
Macaca nemestrina'', ''
Macaca radiata'' and several ''
Presbytis'' species
★ ''P. jefferyi'' - several
gibbon (''
Hylobates'') species
★ ''P. joyeuxi'' - ''
Cercopithecis callitricus''
★ ''P. knowlesi'' - ''
Macaca fascicularis'', ''
Macaca nemestrina'', and ''
Presbytis malalophus''
★ ''P. knowlesi edesoni'' - the
Javanese long-tailed
macaque (''
Macaca irus'')
★ ''P. lemuris'' - the
lemur ''
Lemur collaris'', ''
Lemur macaco macaco''
★ ''P. percygarnhami'' - ''
Lemur macaco macaco''
★ ''P. petersi'' - ''
Cercocebus albigena''
★ ''P. pitheci'' -
orangutans (''
Pongo pygmaeus'')
★ ''P. reichenowi'' - chimpanzee (''
Pan'') species and gorilla (''
Gorilla'') species
★ ''P. rodhaini'' - chimpanzee (''
Pan'') species and gorilla (''
Gorilla'') species
★ ''P. sandoshami'' - the
Malayan flying
lemur (''
Cynocephalus variegatus'')
★ ''P. semnopitheci'' - ''
Semnopithecus entellus''
★ ''P. schwetzi'' - chimpanzee (''
Pan'') species and gorilla (''
Gorilla'') species
★ ''P. semiovale'' - ''
Macaca sinica''
★ ''P. shortii'' - ''
Macaca radiata'', and ''
Macaca sinica''
★ ''P. silvaticum'' -
orangutans (''
Pongo pygmaeus'')
★ ''P. simium'' - the
woolly spider monkey (''
Brachyteles arachnoides'') and several
howler monkeys (''
Alouatta'') species including ''
Alouatta fusca''
★ ''P. uilenbergi'' - ''
Lemur fulvus fulvus''
★ ''P. vivax'' -
orangutans (''
Pongo species'') and
chimpanzees (''
Pan'' species)
★ ''P. youngei'' - the white handed
gibbon (''
Hylobates lar'')
'
Mosquito vectors'
★ ''
Anopheles dirus'' - ''P. cynomolgi'', ''P. inui''
★ ''
Anopheles farauti'' - ''P. coatneyi'', ''P. vivax''
[Collins WE,Sullivan JS,Nace D, Williams T,Sullivan JJ, Galland GG,Grady KK and Bounngaseng A. 2002. Experimental infection of ''Anopheles farauti'' with different species of ''Plasmodium''. J. Parasitol. 88(2):295-298. ]
★ ''
Anopheles funestus'' - ''P. falciparum''
★ ''
Anopheles gambiae'' - ''P. falciparum''
★ ''
Anopheles maculatus'' - ''P. youngei''
★ ''
Anopheles maculipennis'' - ''P. vivax''
★ ''
Anopheles punctipennis'' - ''P. vivax''
★ ''
Anopheles quadrimaculatus'' - ''P. vivax''
★ ''
Anopheles stephensi'' - ''P. cynomogli'', ''P. inui''
★ ''
Anopheles sundaicus'' - ''P. youngei''
★ ''
Anopheles tessellatus'' - ''P. falciparum'', ''P. vivax''
'Subspecies'
★ ''P. cynomolgi'' - ''P. cynomolgi bastianelli'' and ''P. cynomolgi ceylonensis''.
★ ''P. inui'' - ''P. inui inui'' and ''P. inui shortii''
★ ''P. knowlesi'' - ''P. knowlesi edesoni'' and ''P. knowlesi knowlesi''.
★ ''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''
★ ''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'
★ ''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''.
Infections in non primate mammals
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.
'Host records'
★ ''P. aegyptensis'' -
Egyptian grass rat (''
Arvicanthis noloticus'')
[ Abd-el-Aziz GA, Landau I, and Miltgen F. (1975) Description of Plasmodium aegyptensis n. sp., presumed parasite of the Muridae Arvicanthis noloticus in Upper Egypt Ann Parasitol Hum Comp. 50(4):419-424.]
★ ''P. atheruri'' -
African
porcupine (''
Atherurus africanus''), large vesper
mouse (''
Calomys callosus'') and ''
Meriones unguiculatus''
★ ''P. berghei'' - the thicket rat (''
Grammomys surdaster'')
★ ''P. booliati'' -
Malayan giant flying squirrel[ Sandosham AA, Yap LF, Omar I. (1965) A malaria parasite, plasmodium (Vinckeia) booliati sp.nov., from a Malayan giant flying squirrel. Med J Malaya. 20(1):3-7 ]
★ ''P. brodeni'' - elephant shrews (''
Petrodomus teradactylus'')
★ ''P. bubalis'' -
water buffaloes (''
Bubalus bubalis'')
★ ''P. caprae'' - domestic
goat (''
Capra hircus'')
★ ''P. cephalophi'' - the antelope (''
Cephalophus grimmi'') and the grey
duiker (''
Sylvicapra grimmia'')
[bat (''Hipposideros cyclops'')][ Landau I, and Chabaud AG. (1978) Description of P. cyclopsi n. sp. a parasite of the microchiropteran bat Hipposideros cyclops in Gabon. Ann. Parasitol. Hum. Comp. 53(3):247-253.]
★ ''P. odocoilei'' - white tailed deer (''Odocoileus virginianus'')
★ ''P. sandoshami'' - the Sunda flying lemur (''Galeopterus variegatus'')
★ ''P. traguli'' - the mouse deer
★ ''P. tyrio'' - the anteater (''Manus pentadactyla'')
★ ''P. voltaicum'' - the fruit bat (''Roussettus smithi'')
★ ''P. watteni'' - Formosan giant flying squirrel (''Petaurista petaurista grandis'')[Anopheles stephensi'' - ''P. atheruri'', ''P berghei'', ''P. chabaudi'', ''P. yoelii'']
'Subspecies'
★ ''P. berghei'' - ''P. berghei yoelii''
★ ''P. chabaudi'' - ''P. chabaudi adami'' and ''P. chabaudi chabaudi''
★ ''P. melanipherum'' - ''P. melanipherum monosoma''
★ ''P. vinkei'' - ''P. vinckei brucechwatti'', ''P. vinckei petteri'' and ''P. vinckei vinckei''.
★ ''P. yoellii'' - ''P. yoelli nigeriensis'' and ''P. yoelli yoelli''.
'Notes'
★ ''Calomys callosus'' seems unlikely to be a natural host for ''P. atheruri'' as ''P. atheruri'' is found in Africa and ''Calomys callosus'' in South America.
'Less well documented species'
The species listed here from taken from Courtney ''et al''.[12]. should be regarded as dubious.
''P. achromaticum'' - the bat (''Achromaticatus vesperuginis'')
''P. brodini'' - the jumping rat (''Petrodromus tetradactylus'')
''P. melanipherum'' - Schreiber's bat (''Miniopterus schreibersi'')
''P. melanipherum monosoma'' - the bat (''Vesperugo abramus'')
''P. murinum'' - the bat (''Vespertilio murinus'')
'Reclassification'
The literature is replete with species initially classified as ''Plasmodium'' that have been subsequently reclassified. With DNA taxonomy some of these may be once again be classified as ''Plasmodium''. Some of these species are listed here for completness.
''P. epomophori'' of the bat (''Hypsignathus monstruosus'') has been reclassified as ''Hepatocystis epomophori.''
Infections in birds
Species in five of these subgenera infect birds - ''Bennettinia'', ''Giovannolaia'', ''Haemamoeba'', ''Huffia'' and ''Novyella''.[Wiersch SC, Maier WA, Kampen H. Plasmodium (Haemamoeba) cathemerium gene sequences for phylogenetic analysis of malaria parasites. Parasitol Res. 96(2): 90-94 ] ''Giovannolaia'' appears to be a polyphytic group and may be sudivided in the future.[ Martinsen ES,Waite JL,Schall JJ Morphologically defined subgenera of Plasmodium from avian hosts: test of monophyly by phylogenetic analysis of two mitochondrial genes (2006) Parasitology 1 - 8 ]
'Listing'
Species infecting birds include: ''P. ashfordi'', ''P. anasum'', ''P. bambusicolai'', ''P. bigueti'', ''P. biziurae'', ''P. cathemerium'', ''P. circumflexum'', ''P. coggeshalli'', '' P. corradettii'', ''P. coturnix'', ''P. dissanaikei'', ''P. durae'', ''P. elongatum'', ''P. fallax'', ''P forresteri'', ''P. gallinacium'', ''P. garnhami'', ''P. giovannolai'', ''P. griffithsi'', ''P. gundersi'', ''P. guangdong'', ''P. hegneri'', ''P. hermani'', ''P. hexamerium'', ''P. huffi'', ''P. jiangi'', ''P. juxtanucleare'', ''P. kempi'', ''P. lophurae'', ''P.lutzi'', ''P. matutinum'', ''P. nucleophilum'', ''P. papernai'', ''P. paranucleophilum'', ''P. parvulum'', ''P. pediocetti'', ''P. paddae'', ''P. pinotti'', ''P. polare'', ''P. relictum'', ''P. rouxi'', ''P. tenue'', ''P. tejerai'', ''P. tumbayaensis'' and ''P. vaughani''.
'Host records'
★ ''P. ashfordi'' - great reed warblers (''Acrocephalus arundinaceus'')[13]
★ ''P. bigueti'' - the house sparrow (''Passer domesticus'')[14]
★ ''P. biziurae'' - the musk duck (''Biziura lobata'')
★ ''P. cathemerium'' - red-winged blackbird (''Agelaius phoeniceusp''), great horned owl (''Bubo virginianus''), house finch (''Carpodacus mexicanus''), blue jay (''Cyanocitta cristata''), wood thrush (''Hylocichla mustelina''), song sparrow (''Melospiza melodia''), Northern Mockingbird (''Mimus polyglottos leucopterus''), cowbirds (''Molothrus ater ater''), house sparrow (''Passer domesticus''), magpies (''Pica pica budsonia''), bronze grackle (''Quiscalus quiscuIa aeneus'') , finch (''Richmondena cardinalis''), canary (''Serinus canaria''), starling (''Sturnus vulgaris''), house wren (''Troglodytes aedon''), robin (''Turdus migratorius''), white-throated sparrow (''Zonotrichia albicollis'')
★ ''P. circumflexum'' - sharp-shinned hawk, (''Accipiter striatus'')[ Kirkpatrick CE, Lauer DM. (1985) Hematozoa of raptors from southern New Jersey and adjacent areas. J Wildl. Dis. 21(1):1-6. ] helmeted guineafowls, (''Numida meleagris''),[15] red-winged blackbird (''Agelaius phoeniceus''), blue jay (''Cyanocitta cristata''), Cape May warbler (''Dendroica tigrina''), gray cat bird (''Dumella carolinensis''), juncos (''Junco hyemalis byemalls''), song sparrow (''Melospiza melodia''), cowbirds (''Molothrus ater ater'', chestnut-tailed starling (''Sturnus malabaricus''), finch (''Richmondena cardinalis cardinalis''), trumpeter swans (''Olor buccinator''), brown thrasher (''Toxostomar ufum''), robin (''Turdus migratorius''), white-throated sparrow (''Zonotrichia albicollis'')
★ ''P. dissanaikei'' - Ross-ringed parakeet (''Psittacula krameri manillensis'')
★ ''P. durae'' - turkeys (''Meleagris'' species), the common peafowl (''Pavo cristatus''), francolins (''Franoclinus leucoscepus'' and ''Franoclinus levialanti levialanti''), Japanese quail (''Coturnix japonica'') and Lady Amherst pheasents (''Chrysophus amherstiae'')
★ ''P. elongatum'' - red-tailed hawk (''Buteo jamaicensis''), bobwhite quail (''Colinus virginianus virginianus''), bald eagle (''Haliaeetus leucocephalus''), honeycreeper (''Loxops parva''), eastern screech owl (''Otus asio''), black-footed penguins (''Spheniscus demersus''),
★ ''P. fallax'' - pygmy owl (''Glaucidium passerinum''), turkeys (''Meleagris'' species), the helmeted guineafowl (''Numida meleagris'')
★ ''P. forresteri'' - eastern screech-owls (''Otus asio''), great horned owls (''Bubo virginianus''), barred owls (''Strix varia''), bald eagles (''Haliaeetus leucocephalus''), red-shouldered hawks (''Buteo lineatus''), broad-winged hawks (''Buteo platypterus'') and red-tailed hawks (''Buteo jamaicensis'')
★ ''P. gallinaceum'' - red junglefowl (''Gallus gallus'')
★ ''P. garnhami'' - the rain quail (''Coturnix coromendalica'')
★ ''P. giovannolai'' - red-billed choughs (''Pyrrhocorax pyrrhocorax''), the blackbird (''Turdus merula'')
★ ''P. griffithsi'' - wild turkeys (''Meleagris gallopavo intermedia'')
★ ''P. gundersi'' - Owls (''Otus asio'')
★ ''P. guangdong'' - Red-whiskered Bulbul (''Pycnonotus jocosus'')
★ ''P. hegneri'' - common teal (''Anas crecca'')
★ ''P. hermani'' - turkeys (''Meleagris gallopavo'') and bobwhites (''Colinus virginianus'')
★ ''P. hexamerium'' - bluebirds
★ ''P. jiangi'' - the red-whiskered bulbul (''Pycnonotus jocosus'')
★ ''P. juxtanucleare'' - red junglefowl (''Gallus gallus''), black-footed penguins (''Spheniscus demersus'')
★ ''P. kempi'' - turkeys (''Meleagris gallopavo''), bobwhites (''Colinus virginianus''), chukars (''Alectoris graeca''), guinea fowl (''Numida meleagris''), peacocks (''Pavo cristatus'') and canaries (''Serinus canaria''). Mallards (''Anas platyrhynchos'') and domestic geese (''Anser anser'') may be transiently infected.[16]
★ ''P. loprae'' - Peking duck (''Anas platyrhynchos'')
★ ''P. matutinum'' - pigeons and doves (''Colubma'' species)
★ ''P. nucleophilium'' - great tit (''Parus major''), gray catbird (''Dumetella carolinensis'')
★ ''P. nucleophilum toucani'' - Swainson's Toucan (''Ramphastos swainsonii'')
★ ''P. octamerium'' - pintail whydah bird (''Vidua macroura'')[17]
★ ''P. paddae'' - the Java Sparrow (''Padda oryzivora'')
★ ''P. paranucleophilum'' - South American tanager
★ ''P. parvulum'' - vanga species
★ ''P. pedioecetii'' - lesser prairie-chicken (''Tympanuchus pallidicinctus''), Darwin's Nothura (''Nothura darwinii''), grouse
★ ''P. pinotti'' - the Bananaquit (''Coereba flaveola''), ''Euneornis campestris'', ''Loxipasser anoxanthus'', the black-faced Grassquit (''Tiaris bicolor'')
★ ''P. polare'' - Bald Eagle (''Haliaeetus leucocephalus''), the Barn Swallow (''Hirundo rustica''), yellow wagtails (''Motacilla flava'')[18] and cliff swallows (''Petrochelidon pyrrhonota'')
★ ''P. relictum'' - the little night owl (''Athene noctua''), blue quails (''Coturnix chinensis''), Gyr falcons (''Falco rusticolus''), red-backed shrike (''Lanius collurio''), Hawaiian honeycreepers, yellow wagtails (''Motacilla flava''), the house sparrow (''Passer domesticus''), red-billed choughs (''Pyrrhocorax pyrrhocorax''), the tree sparrow (''Passer montanus''), the great tit (''Parus major''), the bearded tit (''Panurus biarmicus''), Magellanic penguins (''Spheniscus magellanicus''), black-footed penguins (''Spheniscus demersus''), pheasents (''Tragopan satyra''), ''Turdus jamaicensis'', the yellow-faced Grassquit (''Tiaris olivacea'')
★ ''P. rouxi'' - partridges
★ ''P. tenue'' - a babbler (''Garrulax canorus taewanus'') and the Pekin Robin (''Leiothrix luteus'')
★ ''P. tejerai'' - domestic turkeys (''Meleagris gallopavo'')
★ ''P. tumbayaensis'' - the thrush (''Planethicus anthracinus'')
★ ''P. vaughani'' - blue jay (''Cyanocitta cristata''), robins (''Erithacus rubecula''), red-billed Leiothrix (''Leiothrix lutea''), ''Loxigilla violacea'', starlings (''Sturnus vulgaris''), juncos (''Junco hyemalis hyemalis''), the house sparrow (''Passer domesticus''), eastern meadowlark (''Sturnella magna''), starling (''Sturnus vulgaris''), Black-faced Grassquit (''Tiaris bicolor'') and ''Turdus jamaicensis''
'Vectors'
★ ''Aedes'' species:
★
★ ''Aedes aegypti'' - ''P. gallinacium''
★ ''Culex'' species:
★
★ ''Culex fatigans'' - ''P. relictum''
★
★ ''Culex pipiens'' - ''P. cathermerium'', ''P. paddae''
★
★ ''Culex pipiens pipiens'' - ''P. kempi''
★
★ ''Culex nigripalpus'' - ''P. elongatum'', ''P. hermani''
★
★ ''Culex quinquefasciatus'' - ''P. relictum''
★
★ ''Culex restuans'' - ''P. elongatum''
★
★ ''Culex salinarius'' - ''P. elongatum'', ''P. hermani''
★
★ ''Culex stigmatastoma'' - ''P. relictum''
★
★ ''Culex tarsalis'' - ''P. kempi'', ''P. hexamerium'', ''P. relictum''
★ ''Mansonia'' species:
★
★ ''Mansionia crassipes'' - ''P. gallinacium''
'Subspecies'
★ ''P. relictum'' has been divded into subspecies: ''P. relictum capistranoae'', ''P. relicturn matutinum'' and ''P. relictum relictum''.
★ ''P. nucleophilum'' has at least one subspecies - ''P. nucleophilum toucani''
'Interelatedness'
★ ''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''.
'Notes'
★ ''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. vaughani'' is the second commonest species of avian malaria parasites after ''P. relictum''.
★ ''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. elongatum'' infects 21 bird families and 59 species of bird. Additional host species are given under the link ''P. elongatum''.
★ ''P. dominicana'' is species known only from fossil amber.[ Poinar, G. (2005) Plasmodium dominicana n. sp. (Plasmodiidae: Haemospororida) from Tertiary Dominican amber. Systematic Parasitology 61 (1) 47-52] It is thought to have been a species infecting birds.
★ The taxonomic status of ''P. corradettii'' (Laird, 1998) is currently regarded as dubious and may be revised.
★ ''P. huffi'' may be the same species as ''P. nucleophilum toucani''.
★ ''P. oti'' is now regarded as the same species as ''P. hexamerium''.
★ 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 recongised 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[19] but this appears to be the only report of this species and it should therefore be regarded of dubious validity.
Infections in reptiles
Species in the subgenera ''Asiamoeba'', ''Carinamoeba'', ''Lacertaemoba'', ''Paraplasmodium'' and ''Sauramoeba'' infect reptiles.[ Schall JJ (2000) Transmission success of the malaria parasite Plasmodium mexicanum into its vector: role of gametocyte density and sex ratio. Parasitology. 121 (6):575-580 ].
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.
'Listing'
Species infecting reptiles include: ''P. achiotense'', ''P. aeuminatum'', ''P. agamae'', ''P. arachniformis'', ''P. attenuatum'',''P. aurulentum'', ''P. australis'', ''P. azurophilum'', ''P. balli'', ''P. basilisci'', ''P. beebei'', ''P. beltrani '', ''P. brumpti'', ''P. brygooi'', ''P. chiricahuae'', ''P. circularis'', ''P. cnemidophori'', ''P. colombiense'', ''P. cordyli'', ''P. diminutivum'', ''P. diploglossi'', ''P. egerniae'', ''P. fairchildi'', ''P. floridense'', ''P. gabaldoni'', ''P. giganteum'', ''P. gologoense'', ''P. gracilis'', ''P. guyannense'', ''P. heischi'', ''P. holaspi'', ''P. icipeensis'', ''P. iguanae'', ''P. josephinae'', ''P. kentropyxi'', ''P. lacertiliae'', ''Plasmodium lainsoni'', ''P. lepidoptiformis'', ''P. lionatum'', ''P. lygosomae'', ''P. mabuiae'', ''P. mackerrasae'', ''P. maculilabre'', ''P. marginatum'', ''P. mexicanum'', ''P. michikoa'', ''P. minasense'', ''P. pelaezi'', ''P. pessoai'', ''P. pifanoi'', ''P. pitmani'', ''P. rhadinurum'', ''P. sasai'',''P. saurocaudatum'', ''P. scorzai'', ''P. siamense'', ''P. robinsoni'', ''P. sasai'', ''P. scorzai'', ''P. tanzaniae'', ''P. tomodoni'', ''P. torrealbai'', ''P. tribolonoti'', ''P. tropiduri'', ''P. uzungwiense'', ''P. vacuolatum'', ''P. vastator'', ''P. volans'', ''P. wenyoni'' and ''P. zonuriae''.
'Host records'
★ ''P. agamae'' - the rainbow lizard (''Agama agama'')
★ ''P. attenuatum'' - ''Ameiva ameiva''
★ ''P. arachniformis'' - chameleons
★ ''P. aurulentum'' - neotropical forest gecko (''Thecadactylus rapicaudus'')
★ ''P. azurophilum'' - anole lizards (''Anolis gingivinus'', ''Anolis gundlachi'', ''Anolis sabanus'')
★ ''P. bailli'' - Anolis lizards (''Anolis limifrons'', ''Anolis lionotus'' and ''Anolis poecilopus'')
★ ''P. basilisci'' - the strpped basilisk (''Basiliscus vittatus'')
★ ''P. beebei'' - the gekko (''Gonatodes taniae'')
★ ''P. brygooi'' - short-horned chameleon (''Chamaeleo brevicornis'')
★ ''P. chiricahuae'' - fence lizard (''Sceloporus jarrovi'')
★ ''P. circularis'' - Australian skink (''Egernia stokesii'')
★ ''P. cnemidophori'' - ''Ameiva ameiva''
★ ''P. colombiense'' - Anolis lizard (''Anolis auratus'')
★ ''P. diploglossi'' - Anguid lizard (''Diploglossus fasciatus'')
★ ''P. draconis'' - the flying lizard (''Draco volans'')
★ ''P. egerniae'' - the land mullet (''Egernia major'')
★ ''P. fairchildi'' - Anolis lizard ''Anolis cupreus''
★ ''P. floridense'' - anole lizards (''Anolis biporcatus'', ''Anolis carolinensis'', ''Anolis frenatus'', ''Anolis gingivinus'', ''Anolis gundlachi'', ''Anolis limifrons'', ''Anolis pentaprion'', ''Anolis sabanus'' and ''Anolis sagrei'')
★ ''P. giganteum'' - the rainbow lizard (''Agama agama''), the African tropical lizard (''Agama cyanogaster'')[ Southgate BA. (1970) Plasmodium (Sauramoeba) giganteum in Agama cyanogaster: a new host record. Trans R Soc Trop Med Hyg. 64(1):12-13 ]
★ ''P. gologoense'' - chameleons
★ ''P. gracilis'' - skink (''Tribolonotus gracilis'')
★ ''P. guyannense'' - the iguanid lizard (''Plica plica'')
★ ''P. heischi'' - skinks (''Mabuya striata'')[ Garnham PC, Telford SR Jr. (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):518-521.]
★ ''P. holaspi'' - African flying lizard (''Holaspis guentheri'')
★ ''P. iguanae'' - ''Iguana iguana iguana''
★ ''P. kentropyxi'' - teiid lizard (''Kentropyx calcarata'')
★ ''P. lacertiliae'' - crocodile skink (''Tribolonotus'' species)
★ ''P. lainsoni'' - the gekko (''Phyllodactylus ventralis'')
★ ''P. lepidoptiformis'' - teiid lizard (''Kentropyx calcarata'')
★ ''P. lionatum'' - the flying gecko (''Ptychozoon lionatum'')
★ ''P. lygosomae'' - skink (''Lygosoma moco'')
★ ''P. mabuiae'' - African skink (''Mabuya quinquetaeiata'')
★ ''P. mackerrasae'' - Australian skinks (''Egernia cunninghami'', ''Egernia stokesii'' and ''Egernia striolata'')
★ ''P. maculilabre'' - African skink (''Mabuya'' species)
★ ''P. marginatum'' - Anolis lizard (''Anolis frenatus'')
★ ''P. mexicanum'' - fence lizards (''Sceloporus occidentalis'')
★ ''P. michikoa'' - chameleons
★ ''P. minasense'' - African skink (''Mabuya agilis'')
★ ''P. minasense anolisi'' - anolis lizards (''Anolis cybotes'', ''Anolis distichus'', ''Anolis frenatus'' and ''Anolis limifrons'')
★ ''P. minasense calcaratae'' - teiid lizard (''Kentropyx calcarata'')
★ ''P. minasense capitoi'' - Anolis lizard (''Anolis capito'')
★ ''P. minasense carinii'' - iguanid lizard (''Iguana iguana'')
★ ''P. minasense diminutivum'' - dwarf tegu lizard (''Ameiva ameiva'')
★ ''P. minasense minasense'' - African skink (''Mabuya mabouya'')
★ ''P. minasense plicae'' - olive tree runner lizard (''Plica umbra'')
★ ''P. minasense tegui'' - blue tengu lizard (''Tupinambis teguixin'')
★ ''P. pelaezi'' - the iguanid lizard (''Urosaurus bicarinatus bicarinatus'')
★ ''P. pessoai'' - snakes (''Spilotes pullatus'' and ''Lachesis muta'')
★ ''P. pifanoi'' - the green ameiva lizard (''Ameiva ameiva'') and a teiid lizard (''Kentropyx calcarata'')
★ ''P. pitmani'' - African skink (''Mabuya'' species)
★ ''P. rhadinurum'' - Iguana iguana iguana
★ ''P. robinsoni'' - the Parson's Chameleon (''Chamaelo parsoni crucifer'')
★ ''P. sasai'' - Japanese lacertids (''Takydromus tachydromoides'' and ''Takydromus smaragdinus'')
★ ''P. saurocaudatum'' - the many-lined sun skink (''Mabuya multifasciata]]'')
★ ''P. scorzai'' - the gekko ''Phyllodactylus ventralis''
★ ''P. siamense'' - lizards.[ Telford SR (1986) Fallisia parasites (Haemosporidia: Plasmodiidae) from the flying lizard, Draco maculatus (Agamidae) in Thailand. J Parasitol. 72(5):766-769 ]
★ ''P. tanzaniae'' - chameleons
★ ''P. tomodoni'' - snakes
★ ''P. torrealbai'' - Anolis lizards
★ ''P. tribolonoti'' - skinks (''Tribolonotus gracilis'')
★ ''P. tropiduri'' - iguanid lizard (''Tropidurus torquatus''), Anolis lizards (''Anolis biporcatus'',''Anolis cybotes'', ''Anolis frenatus'', ''Anolis limifrons'', ''Anolis lionotus'', ''Anolis pentaprion'' and ''Anolis poecilopus''), teiid lizard (''Kentropyx calcarata'')
★ ''P. tropiduri aquaticum'' - Anolis lizards (''Anolis lionotus'' and ''Anolis poecilopus'')
★ ''P. tropiduri panamense'' - Anolis lizard (''Anolis biporcatus'')
★ ''P. tropiduri tropiduri'' - ''Tropidurus hispidus''
★ ''P. robinsoni'' - the red-headed rock agama lizard (''Agama agama'')
★ ''P. uzungwiense'' - chameleons
★ ''P. vacuolatum'' - the rainforest lizard (''Plica umbra'')
★ ''P. vastator'' - the flying lizard (''Draco volans'')
★ ''P. volans'' - the flying lizard (''Draco volans'')
★ ''P. wenyoni'' - snakes
'Vectors'
★ ''P. agamae'' - ''Lutzomyia'' or ''Culicoides'' species
Culex:
★ ''Culex fatigans'' - ''P. rhadinurum''
Aedes:
★ ''Aedes aegypti'' - ''P. rhadinurum''
'Subspecies'
★ ''P. fairchildi'' - ''P. fairchildi fairchildi'' and ''P. fairchildi hispaniolae''
★ ''P. lygosomae'' - ''P. lygosomae nucleoversans'' and ''P. lygosomae nucleoversans''
★ ''P. minasense'' - ''P. minasense anolisi'', ''P. minasense capitoi'', ''P. minasense carinii'', ''P. minasense diminutivum'', ''P. minasense minasense'', ''P. minasense plicae'', and ''P. minasense tegui''.[ Telford SR Jr. (1979) A taxonomic revision of small neotropical saurian Malarias allied to Plasmodium minasense. Ann Parasitol Hum Comp. 54(4):409-422] An additional subspecies ''P. minasense calcaratae'' has also been described.[Telford SR Jr and Telford SR 3rd. Rediscovery and redescription of Plasmodium pifanoi and description of two additional Plasmodium parasites of Venezuelan lizards. Journal of Parasitology (2003) 89(2):362-368]
★ ''P. traguli'' - ''P. traguli traguli'' and ''P. traguli memmina''.
★ ''P. tropiduri'' - ''P. tropiduri aquaticum'', ''P. tropiduri panamense'' and ''P. tropiduri tropiduri''.[ Telford SR Jr.A taxonomic reconsideration of some Plasmodium species from iguanid lizards. Ann Parasitol Hum Comp. (1979) 54(2):129-144 ]
'Interrelatedness'
★ ''P. floridense'' is closely related to ''P. tropiduri'' and ''P. minasense''
Species reclassified into other genera
The following species are currently (2007) regarded as belonging to the genus ''Hepatocystis'' rather than ''Plasmodium''.
★ ''Plasmodium epomophori''
★ ''Plasmodium kochi''
★ ''Plasmodium limnotragi'' Van Denberghe 1937
★ ''Plasmodium pteropi'' Breinl 1911
★ ''Plasmodium ratufae'' Donavan 1920
★ ''Plasmodium vassali'' Laveran 1905
''Plasmodium gonatodi'' has been reclassified as a species of ''Garnia'' and has been renamed ''Garnia gonatodi''.
References
1. Chavatte JM, Chiron F, Chabaud A, Landau I. (2007) Probable speciations by "host-vector 'fidelisation'": 14 species of Plasmodium from Magpies. Parasite 14(1):21-37
2. [http://www.hhmi.org/news/menard20060122.html HHMI Staff (22 January 2006) "Malaria Parasites Develop in Lymph Nodes" ''HHMI News'' Howard Hughes Medical Institute
3. Corradetti A., Garnham P. C. C. and Laird M. (1963). New classification of the avian malaria parasites. Parassitologia 5, 1–4
4. Valkiunas, G. (1997). Bird Haemosporidia. Institute of Ecology, Vilnius
5. Martinsen,E. S., Waite J. L. and Schall J. J. (2006) Morphologically defined subgenera of Plasmodium from avian hosts: test of monophyly by phylogenetic analysis of two mitochondrial genes. Parasitology 1-8
6. Garnham P.C.C. (1966) Malaria parasites and other haemospordia. Oxford, Blackwell
7. Telford S. (1988) A contribution to the systematics of the reptilian malaria parasites, family Plasmodiidae (Apicomplexa: Haemosporina). Bulletin of the Florida State Museum Biological Sciences 34, 65-96
8. 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
9. Russel P.F. (1928) ''Plasmodium tenue'' (Stephens): A review of the literature and a case report. Am. J. Trop. Med. s1-8(5) 449-479
10. Reid MJ. (2006) Transmission of Human and Macaque Plasmodium spp. to Ex-Captive Orangutans in Kalimantan,
Indonesia. Emerg Infect Dis. 12(12):1902-1908
11. Coatney G. R. and Roudabush R. L. (1936) A catalog and host-index of the genus Plasmodium. J. Parasitol. 22 (4) 338-353
12. Coatney G. R. and Roudabush R. L. (1936) A catalog and host-index of the genus ''Plasmodium''. J. Parasitol. 22 (4) 338-353
13. Valkiūnas G., Zehtindjiev P., Hellgren O., Ilieva M., Iezhova T. A. and 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.
14. 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.
15. 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.
16. Christensen BM, Barnes HJ, Rowley WA. (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
17. Manwell RD. (1968) Plasmodium octamerium n. sp., an avian malaria parasite from the pintail whydah bird Vidua macroura. J. Protozool. 15(4):680-685
18. Valkiunas G, Iezhova TA. (2001) A comparison of the blood parasites in three subspecies of the yellow wagtail Motacilla flava. J. Parasitol. 87(4):930-934.
19. Manwell RD (1966) Plasmodium japonicum, P. juxtanucleare and P. nucleophilum in the Far East. J. Protozool. 13(1):8-11.
General references
The standard reference books for the identification of ''Plasmodium'' species are:
★ Laird, M. (1998) Avian Malaria in the Asian Tropical Subregion. Springer, Singapore.
★ Garnham P.C (1966) Malaria Parasites And Other Haemosporidia. Blackwell, Oxford. This book remains the standard reference work on malarial species classification.
★ Hewitt (1940) Bird Malaria. Baltimore, The Johns Hopkins Press.
Other useful references include
★ Short, H. E. (1951) ''Life-cycle of the mammalian malaria parasite" ''British Medical Bulletin'' 8(1): pp. 7-9, (PMID 14944807);
★ Baldacci, Patricia and Ménard, Robert (Oct. 2004) "The elusive malaria sporozoite in the mammalian host" ''Molecular Microbiology'' 54(2): pp. 298-306, (AN 14621725);
★ Bledsoe, G. H. (December 2005) "Malaria primer for clinicians in the United States" ''Southern Medical Journal'' 98(12): pp. 1197-204 (PMID 16440920);
Some history of malaria - http://muse.jhu.edu/journals/bulletin_of_the_history_of_medicine/v079/79.2slater.html
External links
★
★ Malaria Atlas Project
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