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dium belongs to the phylum apicomplexa a taxonomic group of single celled parasites with characteristic secretory organelles at one end of the cell 15 within apicomplexa plasmodium is within the order haemosporida a group that includes all apicomplexans that live within blood cells 16 based on the presence of the pigment hemozoin and the method of asexual reproduction the order is further split into four families of which plasmodium is in the family plasmodiidae 17 the genus plasmodium consists of over 200 species generally described on the basis of their appearance in blood smears of infected vertebrates 18 these species have been categorized on the basis of their morphology and host range into 14 subgenera 17 subgenus asiamoeba telford 1988 reptiles subgenus bennettinia valkiunas 1997 birds subgenus carinamoeba garnham 1966 reptiles subgenus giovannolaia corradetti et al 1963 birds subgenus haemamoeba corradetti et al 1963 birds subgenus huffia corradetti et al 1963 birds subgenus lacertamoeba telford 1988 reptiles subgenus laverania bray 1958 great apes humans subgenus novyella corradetti et al 1963 birds subgenus ophidiella telford 1988 reptiles subgenus paraplasmodium telford 1988 reptiles subgenus plasmodium bray 1955 monkeys and apes subgenus sauramoeba garnham 1966 reptiles subgenus vinckeia garnham 1964 mammals inc primates species infecting monkeys and apes with the exceptions of p falciparum and p reichenowi which together make up the subgenus laverania are classified in the subgenus plasmodium parasites infecting other mammals including some primates lemurs and others are classified in the subgenus vinckeia the five subgenera bennettinia giovannolaia haemamoeba huffia and novyella contain the known avian malarial species 19 the remaining subgenera asiamoeba carinamoeba lacertamoeba ophidiella paraplasmodium and sauramoeba contain the diverse groups of parasites found to infect reptiles 20 phylogeny edit more recent studies of plasmodium species using molecular methods have implied that the group s evolution has not perfectly followed taxonomy 2 many plasmodium species that are morphologically similar or infect the same hosts turn out to be only distantly related 21 in the 1990s several studies sought to evaluate evolutionary relationships of plasmodium species by comparing ribosomal rna and a surface protein gene from various species finding the human parasite p falciparum to be more closely related to avian parasites than to other parasites of primates 17 however later studies sampling more plasmodium species found the parasites of mammals to form a clade along with the genus hepatocystis while the parasites of birds or lizards appear to form a separate clade with evolutionary relationships not following the subgenera 17 22 leucocytozoon haemoproteus plasmodium plasmodium of lizards and birds subgenus laverania subgenus plasmodium subgenus vinckeia hepatocystis parasites of bats estimates for when different plasmodium lineages diverged have differed broadly estimates for the diversification of the order haemosporida range from around 16 2 million to 100 million years ago 17 there has been particular interest in dating the divergence of the human parasite p falciparum from other plasmodium lineages due to its medical importance for this estimated dates range from 110 000 to 2 5 million years ago 17 distribution edit plasmodium species are distributed globally all plasmodium species are parasitic and must pass between a vertebrate host and an insect host to complete their life cycles different species of plasmodium display different host ranges with some species restricted to a single vertebrate and insect host while other species can infect several species of vertebrates and or insects vertebrates edit relative incidence of plasmodium species by country of origin for imported cases to non endemic countries 23 plasmodium parasites have been described in a broad array of vertebrate hosts including reptiles birds and mammals 24 while many species can infect more than one vertebrate host they are generally specific to one of these classes such as birds 24 humans are primarily infected by five species of plasmodium with the overwhelming majority of severe disease and death caused by plasmodium falciparum 25 some species that infect humans can also infect other primates and zoonoses of certain species e g p knowlesi from other primates to humans are common 25 non human primates also contain a variety of plasmodium species that do not generally infect humans some of these can cause severe disease in primates while others can remain in the host for prolonged periods without causing disease 26 many other mammals also carry plasmodium species such as a variety of rodents ungulates and bats again some species of plasmodium can cause severe disease in some of these hosts while many appear not to 27 over 150 species of plasmodium infect a broad variety of birds in general each species of plasmodium infects one to a few species of birds 28 plasmodium parasites that infect birds tend to persist in a given host for years or for the life time of the host although in some cases plasmodium infections can result in severe illness and rapid death 29 30 unlike with plasmodium species infecting mammals those infecting birds are distributed across the globe 28 species from several subgenera of plasmodium infect diverse reptiles plasmodium parasites have been described in most lizard families and like avian parasites are spread worldwide 31 again parasites can result either in severe disease or be apparently asymptomatic depending on the parasite and the host 31 a number of drugs have been developed over the years to control plasmodium infection in vertebrate hosts particularly in humans quinine was used as a frontline antimalarial from the 17th century until widespread resistance emerged in the early 20th century 32 resistance to quinine spurred the development of a broad array of antimalarial medications through the 20th century including chloroquine proguanil atovaquone sulfadoxine pyrimethamine mefloquine and artemisinin 32 in all cases parasites resistant to a given drug have emerged within a few decades of the drugs deployment 32 to combat this antimalarial drugs are frequently used in combination with artemisinin combination therapies currently the gold standard for treatment 33 in general antimalarial drugs target the life stages of plasmodium parasites that reside within vertebrate red blood cells as these are the stages that tend to cause disease 34 however drugs targeting other stages of the parasite life cycle are under development in order to prevent infection in travelers and to prevent transmission of sexual stages to insect hosts 35 a clinic for treating human malaria in tanzania over 3000 species of lizard including the carolina anole anolis carolinensis carry some 90 kinds of malaria insects edit the mosquito anopheles stephensi is among the blood feeding insects that can be infected by a species of plasmodium in addition to a vertebrate host all plasmodium species also infect a bloodsucking insect host generally a mosquito although some reptile infecting parasites are transmitted by sandflies mosquitoes of the genera culex anopheles culiseta mansonia and aedes act as insect hosts for various plasmodium species the best studied of these are the anopheles mosquitoes which host the plasmodium parasites of human malaria as well as culex mosquitoes which host the plasmodium species that cause malaria in birds only female mosquitoes are infected with plasmodium since only they feed on the blood of vertebrate hosts 36 different species affect their insect hosts differently sometimes insects infected with plasmodium have reduced lifespan and reduced ability to produce offspring 37 further some species of plasmodium appear to cause insects to prefer to bite infected vertebrate hosts over non infected hosts 37 38 39 history edit charles louis alphonse laveran first described parasites in the blood of malaria patients in 1880 40 he named the parasite oscillaria malariae 40 in 1885 zoologists ettore marchiafva and angelo celli reexamined the parasite and termed it a member of a new genus plasmodium named for the resemblance to the multinucleate cells of slime molds of the same name 41 notes 1 the fact that several species may be involved in causing different forms of malaria was first recognized by camillo golgi in 1886 40 in 1897 william welch identified and named plasmodium falciparum soon thereafter giovanni batista grassi and raimondo filetti named the parasites causing two different types of human malaria plasmodium vivax and plasmodium malariae 40 this was followed by the recognition of the other two species of plasmodium which infect humans plasmodium ovale 1922 and plasmodium knowlesi identified in long tailed macaques in 1931 in humans in 1965 40 the contribution of insect hosts to the plasmodium life cycle was described in 1897 by ronald ross and in 1899 by giovanni batista grassi amico bignami and giuseppe bastianelli 40 in 1966 cyril garnham proposed separating plasmodium into nine subgenera based on host specificity and parasite morphology 18 this included four subgenera that had previously been proposed for bird infecting plasmodium species by a corradetti in 1963 42 19 this scheme was expanded upon by sam r telford in 1988 when he reclassified plasmodium parasites that infect reptiles adding five subgenera 20 18 in 1997 g valkiunas reclassified the bird infecting plasmodium species adding a fifth subgenus bennettinia 19 43 see also edit haematozoa list of plasmodium species plasmodium molecular tools notes edit the plural of plasmodium is not plasmodia instead multiple species of the genus are referred to as plasmodium species 41 references edit cdc malaria parasites about cdc malaria u s centers for disease control and prevention retrieved 28 december 2015 a b zilversmit m perkins s plasmodium tree of life web project retrieved 1 june 2016 obado samson o glover lucy deitsch kirk w 2016 the nuclear envelope and gene organization in parasitic protozoa specializations associated with disease molecular and biochemical parasitology 209 1 2 104 113 doi 10 1016 j molbiopara 2016 07 008 pmid 27475118 a b jimenez ruiz elena morlon guyot juliette daher wassim meissner markus 2016 vacuolar protein sorting mechanisms in apicomplexan parasites molecular and biochemical parasitology 209 1 2 18 25 doi 10 1016 j molbiopara 2016 01 007 pmc 5154328 pmid 26844642 counihan natalie a kalanon ming coppel ross l de koning ward tania f 2013 plasmodium rhoptry proteins why order is important trends in parasitology 29 5 228 36 doi 10 1016 j pt 2013 03 003 pmid 23570755 a b kemp louise e yamamoto masahiro soldati favre dominique 2013 subversion of host cellular functions by the apicomplexan parasites fems microbiology reviews 37 4 607 31 doi 10 1111 1574 6976 12013 pmid 23186105 a 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