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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Chimpanzee Malaria Parasites Related to <italic>Plasmodium ovale</italic> in Africa</title><author><name sortKey="Duval, Linda" sort="Duval, Linda" uniqKey="Duval L" first="Linda" last="Duval">Linda Duval</name><affiliation><nlm:aff id="aff1"><addr-line>Laboratoire de Biologie fonctionnelle des protozoaires, USM 504, Muséum National d'Histoire Naturelle, Paris, France</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff2"><addr-line>Laboratoire de Pathogénie virale, Institut Pasteur, Paris, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="Nerrienet, Eric" sort="Nerrienet, Eric" uniqKey="Nerrienet E" first="Eric" last="Nerrienet">Eric Nerrienet</name><affiliation><nlm:aff id="aff3"><addr-line>Laboratoire HIV et Hepatites, Institut Pasteur du Cambodge, Phnom Penh, Cambodia</addr-line></nlm:aff></affiliation></author><author><name sortKey="Rousset, Dominique" sort="Rousset, Dominique" uniqKey="Rousset D" first="Dominique" last="Rousset">Dominique Rousset</name><affiliation><nlm:aff id="aff4"><addr-line>Unité de virologie, Centre Pasteur du Cameroun, Yaoundé, Cameroun</addr-line></nlm:aff></affiliation></author><author><name sortKey="Sadeuh Mba, Serge Alain" sort="Sadeuh Mba, Serge Alain" uniqKey="Sadeuh Mba S" first="Serge Alain" last="Sadeuh Mba">Serge Alain Sadeuh Mba</name><affiliation><nlm:aff id="aff4"><addr-line>Unité de virologie, Centre Pasteur du Cameroun, Yaoundé, Cameroun</addr-line></nlm:aff></affiliation></author><author><name sortKey="Houze, Sandrine" sort="Houze, Sandrine" uniqKey="Houze S" first="Sandrine" last="Houze">Sandrine Houze</name><affiliation><nlm:aff id="aff5"><addr-line>Centre National de Référence du Paludisme, AP-HP, Hôpital Bichat-Claude Bernard, Paris, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="Fourment, Mathieu" sort="Fourment, Mathieu" uniqKey="Fourment M" first="Mathieu" last="Fourment">Mathieu Fourment</name><affiliation><nlm:aff id="aff6"><addr-line>Unité de Virologie, Institut Pasteur du Cambodge, Phnom Penh, Cambodia</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff7"><addr-line>Department of Biological Sciences, Macquarie University, Sydney, Australia</addr-line></nlm:aff></affiliation></author><author><name sortKey="Le Bras, Jacques" sort="Le Bras, Jacques" uniqKey="Le Bras J" first="Jacques" last="Le Bras">Jacques Le Bras</name><affiliation><nlm:aff id="aff5"><addr-line>Centre National de Référence du Paludisme, AP-HP, Hôpital Bichat-Claude Bernard, Paris, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="Robert, Vincent" sort="Robert, Vincent" uniqKey="Robert V" first="Vincent" last="Robert">Vincent Robert</name><affiliation><nlm:aff id="aff1"><addr-line>Laboratoire de Biologie fonctionnelle des protozoaires, USM 504, Muséum National d'Histoire Naturelle, Paris, France</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff8"><addr-line>Unité de Recherche Caractérisation et contrôle des populations de vecteurs, UR 16, Institut de Recherche pour le Développement, Montpellier, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="Ariey, Frederic" sort="Ariey, Frederic" uniqKey="Ariey F" first="Frederic" last="Ariey">Frederic Ariey</name><affiliation><nlm:aff id="aff9"><addr-line>Unité d'Epidémiologie Moleculaire, Institut Pasteur du Cambodge, Phnom Penh, Cambodia</addr-line></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">19436742</idno><idno type="pmc">2677663</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2677663</idno><idno type="RBID">PMC:2677663</idno><idno type="doi">10.1371/journal.pone.0005520</idno><date when="2009">2009</date><idno type="wicri:Area/Pmc/Corpus">001526</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">001526</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Chimpanzee Malaria Parasites Related to <italic>Plasmodium ovale</italic> in Africa</title><author><name sortKey="Duval, Linda" sort="Duval, Linda" uniqKey="Duval L" first="Linda" last="Duval">Linda Duval</name><affiliation><nlm:aff id="aff1"><addr-line>Laboratoire de Biologie fonctionnelle des protozoaires, USM 504, Muséum National d'Histoire Naturelle, Paris, France</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff2"><addr-line>Laboratoire de Pathogénie virale, Institut Pasteur, Paris, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="Nerrienet, Eric" sort="Nerrienet, Eric" uniqKey="Nerrienet E" first="Eric" last="Nerrienet">Eric Nerrienet</name><affiliation><nlm:aff id="aff3"><addr-line>Laboratoire HIV et Hepatites, Institut Pasteur du Cambodge, Phnom Penh, Cambodia</addr-line></nlm:aff></affiliation></author><author><name sortKey="Rousset, Dominique" sort="Rousset, Dominique" uniqKey="Rousset D" first="Dominique" last="Rousset">Dominique Rousset</name><affiliation><nlm:aff id="aff4"><addr-line>Unité de virologie, Centre Pasteur du Cameroun, Yaoundé, Cameroun</addr-line></nlm:aff></affiliation></author><author><name sortKey="Sadeuh Mba, Serge Alain" sort="Sadeuh Mba, Serge Alain" uniqKey="Sadeuh Mba S" first="Serge Alain" last="Sadeuh Mba">Serge Alain Sadeuh Mba</name><affiliation><nlm:aff id="aff4"><addr-line>Unité de virologie, Centre Pasteur du Cameroun, Yaoundé, Cameroun</addr-line></nlm:aff></affiliation></author><author><name sortKey="Houze, Sandrine" sort="Houze, Sandrine" uniqKey="Houze S" first="Sandrine" last="Houze">Sandrine Houze</name><affiliation><nlm:aff id="aff5"><addr-line>Centre National de Référence du Paludisme, AP-HP, Hôpital Bichat-Claude Bernard, Paris, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="Fourment, Mathieu" sort="Fourment, Mathieu" uniqKey="Fourment M" first="Mathieu" last="Fourment">Mathieu Fourment</name><affiliation><nlm:aff id="aff6"><addr-line>Unité de Virologie, Institut Pasteur du Cambodge, Phnom Penh, Cambodia</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff7"><addr-line>Department of Biological Sciences, Macquarie University, Sydney, Australia</addr-line></nlm:aff></affiliation></author><author><name sortKey="Le Bras, Jacques" sort="Le Bras, Jacques" uniqKey="Le Bras J" first="Jacques" last="Le Bras">Jacques Le Bras</name><affiliation><nlm:aff id="aff5"><addr-line>Centre National de Référence du Paludisme, AP-HP, Hôpital Bichat-Claude Bernard, Paris, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="Robert, Vincent" sort="Robert, Vincent" uniqKey="Robert V" first="Vincent" last="Robert">Vincent Robert</name><affiliation><nlm:aff id="aff1"><addr-line>Laboratoire de Biologie fonctionnelle des protozoaires, USM 504, Muséum National d'Histoire Naturelle, Paris, France</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff8"><addr-line>Unité de Recherche Caractérisation et contrôle des populations de vecteurs, UR 16, Institut de Recherche pour le Développement, Montpellier, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="Ariey, Frederic" sort="Ariey, Frederic" uniqKey="Ariey F" first="Frederic" last="Ariey">Frederic Ariey</name><affiliation><nlm:aff id="aff9"><addr-line>Unité d'Epidémiologie Moleculaire, Institut Pasteur du Cambodge, Phnom Penh, Cambodia</addr-line></nlm:aff></affiliation></author></analytic><series><title level="j">PLoS ONE</title><idno type="eISSN">1932-6203</idno><imprint><date when="2009">2009</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Since the 1970's, the diversity of <italic>Plasmodium</italic> parasites in African great apes has been neglected. Surprisingly, <italic>P. reichenowi</italic>, a chimpanzee parasite, is the only such parasite to have been molecularly characterized. This parasite is closely phylogenetically related to <italic>P. falciparum</italic>, the principal cause of the greatest malaria burden in humans. Studies of malaria parasites from anthropoid primates may provide relevant phylogenetic information, improving our understanding of the origin and evolutionary history of human malaria species. In this study, we screened 130 DNA samples from chimpanzees (<italic>Pan troglodytes</italic>) and gorillas (<italic>Gorilla gorilla</italic>) from Cameroon for <italic>Plasmodium</italic> infection, using <italic>cytochrome b</italic> molecular tools. Two chimpanzees from the subspecies <italic>Pan t</italic>. <italic>troglodytes</italic> presented single infections with <italic>Plasmodium</italic> strains molecularly related to the human malaria parasite <italic>P. ovale</italic>. These chimpanzee parasites and 13 human strains of <italic>P. ovale</italic> originated from a various sites in Africa and Asia were characterized using <italic>cytochrome b</italic> and <italic>cytochrome c oxidase 1</italic> mitochondrial partial genes and nuclear <italic>ldh</italic> partial gene. Consistent with previous findings, two genetically distinct types of <italic>P. ovale</italic>, classical and variant, were observed in the human population from a variety of geographical locations. One chimpanzee <italic>Plasmodium</italic> strain was genetically identical, on all three markers tested, to variant <italic>P. ovale</italic> type. The other chimpanzee <italic>Plasmodium</italic> strain was different from <italic>P. ovale</italic> strains isolated from humans. This study provides the first evidence of possibility of natural cross-species exchange of <italic>P. ovale</italic> between humans and chimpanzees of the subspecies <italic>Pan t. troglodytes</italic>.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article" xml:lang="EN"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">PLoS ONE</journal-id><journal-id journal-id-type="publisher-id">plos</journal-id><journal-id journal-id-type="pmc">plosone</journal-id><journal-title>PLoS ONE</journal-title><issn pub-type="epub">1932-6203</issn><publisher><publisher-name>Public Library of Science</publisher-name><publisher-loc>San Francisco, USA</publisher-loc></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">19436742</article-id><article-id pub-id-type="pmc">2677663</article-id><article-id pub-id-type="publisher-id">08-PONE-RA-07611R2</article-id><article-id pub-id-type="doi">10.1371/journal.pone.0005520</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group><subj-group subj-group-type="Discipline"><subject>Evolutionary Biology</subject><subject>Infectious Diseases</subject><subject>Microbiology/Parasitology</subject><subject>Molecular Biology/Molecular Evolution</subject></subj-group></article-categories><title-group><article-title>Chimpanzee Malaria Parasites Related to <italic>Plasmodium ovale</italic> in Africa</article-title><alt-title alt-title-type="running-head"><italic>Plasmodium ovale</italic></alt-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Duval</surname><given-names>Linda</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><xref ref-type="aff" rid="aff2"><sup>2</sup></xref><xref ref-type="corresp" rid="cor1"><sup>*</sup></xref></contrib><contrib contrib-type="author"><name><surname>Nerrienet</surname><given-names>Eric</given-names></name><xref ref-type="aff" rid="aff3"><sup>3</sup></xref></contrib><contrib contrib-type="author"><name><surname>Rousset</surname><given-names>Dominique</given-names></name><xref ref-type="aff" rid="aff4"><sup>4</sup></xref></contrib><contrib contrib-type="author"><name><surname>Sadeuh Mba</surname><given-names>Serge Alain</given-names></name><xref ref-type="aff" rid="aff4"><sup>4</sup></xref></contrib><contrib contrib-type="author"><name><surname>Houze</surname><given-names>Sandrine</given-names></name><xref ref-type="aff" rid="aff5"><sup>5</sup></xref></contrib><contrib contrib-type="author"><name><surname>Fourment</surname><given-names>Mathieu</given-names></name><xref ref-type="aff" rid="aff6"><sup>6</sup></xref><xref ref-type="aff" rid="aff7"><sup>7</sup></xref></contrib><contrib contrib-type="author"><name><surname>Le Bras</surname><given-names>Jacques</given-names></name><xref ref-type="aff" rid="aff5"><sup>5</sup></xref></contrib><contrib contrib-type="author"><name><surname>Robert</surname><given-names>Vincent</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><xref ref-type="aff" rid="aff8"><sup>8</sup></xref></contrib><contrib contrib-type="author"><name><surname>Ariey</surname><given-names>Frederic</given-names></name><xref ref-type="aff" rid="aff9"><sup>9</sup></xref></contrib></contrib-group><aff id="aff1"><label>1</label><addr-line>Laboratoire de Biologie fonctionnelle des protozoaires, USM 504, Muséum National d'Histoire Naturelle, Paris, France</addr-line></aff><aff id="aff2"><label>2</label><addr-line>Laboratoire de Pathogénie virale, Institut Pasteur, Paris, France</addr-line></aff><aff id="aff3"><label>3</label><addr-line>Laboratoire HIV et Hepatites, Institut Pasteur du Cambodge, Phnom Penh, Cambodia</addr-line></aff><aff id="aff4"><label>4</label><addr-line>Unité de virologie, Centre Pasteur du Cameroun, Yaoundé, Cameroun</addr-line></aff><aff id="aff5"><label>5</label><addr-line>Centre National de Référence du Paludisme, AP-HP, Hôpital Bichat-Claude Bernard, Paris, France</addr-line></aff><aff id="aff6"><label>6</label><addr-line>Unité de Virologie, Institut Pasteur du Cambodge, Phnom Penh, Cambodia</addr-line></aff><aff id="aff7"><label>7</label><addr-line>Department of Biological Sciences, Macquarie University, Sydney, Australia</addr-line></aff><aff id="aff8"><label>8</label><addr-line>Unité de Recherche Caractérisation et contrôle des populations de vecteurs, UR 16, Institut de Recherche pour le Développement, Montpellier, France</addr-line></aff><aff id="aff9"><label>9</label><addr-line>Unité d'Epidémiologie Moleculaire, Institut Pasteur du Cambodge, Phnom Penh, Cambodia</addr-line></aff><contrib-group><contrib contrib-type="editor"><name><surname>Snounou</surname><given-names>Georges</given-names></name><role>Editor</role><xref ref-type="aff" rid="edit1"></xref></contrib></contrib-group><aff id="edit1">Université Pierre et Marie Curie, France</aff><author-notes><corresp id="cor1">* E-mail: <email>lduval@pasteur.fr</email></corresp><fn fn-type="con"><p>Conceived and designed the experiments: LD EN VR FA. Performed the experiments: LD DR SASM SH. Analyzed the data: LD MF FA. Contributed reagents/materials/analysis tools: LD EN DR SH MF JLB FA. Wrote the paper: LD EN FA. Reviewed the paper: DR SASM SH MF JLB VR.</p></fn></author-notes><pub-date pub-type="collection"><year>2009</year></pub-date><pub-date pub-type="epub"><day>13</day><month>5</month><year>2009</year></pub-date><volume>4</volume><issue>5</issue><elocation-id>e5520</elocation-id><history><date date-type="received"><day>4</day><month>12</month><year>2008</year></date><date date-type="accepted"><day>5</day><month>4</month><year>2009</year></date></history><copyright-statement>Duval et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</copyright-statement><copyright-year>2009</copyright-year><abstract><p>Since the 1970's, the diversity of <italic>Plasmodium</italic> parasites in African great apes has been neglected. Surprisingly, <italic>P. reichenowi</italic>, a chimpanzee parasite, is the only such parasite to have been molecularly characterized. This parasite is closely phylogenetically related to <italic>P. falciparum</italic>, the principal cause of the greatest malaria burden in humans. Studies of malaria parasites from anthropoid primates may provide relevant phylogenetic information, improving our understanding of the origin and evolutionary history of human malaria species. In this study, we screened 130 DNA samples from chimpanzees (<italic>Pan troglodytes</italic>) and gorillas (<italic>Gorilla gorilla</italic>) from Cameroon for <italic>Plasmodium</italic> infection, using <italic>cytochrome b</italic> molecular tools. Two chimpanzees from the subspecies <italic>Pan t</italic>. <italic>troglodytes</italic> presented single infections with <italic>Plasmodium</italic> strains molecularly related to the human malaria parasite <italic>P. ovale</italic>. These chimpanzee parasites and 13 human strains of <italic>P. ovale</italic> originated from a various sites in Africa and Asia were characterized using <italic>cytochrome b</italic> and <italic>cytochrome c oxidase 1</italic> mitochondrial partial genes and nuclear <italic>ldh</italic> partial gene. Consistent with previous findings, two genetically distinct types of <italic>P. ovale</italic>, classical and variant, were observed in the human population from a variety of geographical locations. One chimpanzee <italic>Plasmodium</italic> strain was genetically identical, on all three markers tested, to variant <italic>P. ovale</italic> type. The other chimpanzee <italic>Plasmodium</italic> strain was different from <italic>P. ovale</italic> strains isolated from humans. This study provides the first evidence of possibility of natural cross-species exchange of <italic>P. ovale</italic> between humans and chimpanzees of the subspecies <italic>Pan t. troglodytes</italic>.</p></abstract><counts><page-count count="7"></page-count></counts></article-meta></front><body><sec id="s1"><title>Introduction</title><p><italic>Plasmodium ovale</italic>, <italic>P. falciparum</italic>, <italic>P. vivax</italic> and <italic>P. malariae</italic> belong to phylum Apicomplexa, order Haemosporidia and family Plasmodiidae. Haemosporidia are intracellular parasites transmitted by haematophagous dipterans. They infect a large variety of vertebrate amniotes, such as mammals (including humans), birds, chelonians, squamates, and crocodilians, <xref ref-type="bibr" rid="pone.0005520-Garnham1">[1]</xref>. Some are highly pathogenic and may have important implications for human public health, domestic animal health and wildlife biodiversity conservation <xref ref-type="bibr" rid="pone.0005520-Guerra1">[2]</xref>, <xref ref-type="bibr" rid="pone.0005520-vanRiper1">[3]</xref>.</p><p><italic>P. ovale</italic>, the last of the human malaria parasites to be identified, was described in the blood of an East African patient, by Stephens in 1922. It is a relapse parasite, generating secondary infections that are usually asymptomatic <xref ref-type="bibr" rid="pone.0005520-Collins1">[4]</xref>. However, <italic>P. ovale</italic> may interact with other species of <italic>Plasmodium</italic> infecting humans, such as <italic>P. falciparum</italic> and <italic>P. vivax</italic>, and may have a major influence on the epidemiological features of malaria <xref ref-type="bibr" rid="pone.0005520-Mueller1">[5]</xref>.</p><p>Few epidemiological data are available for <italic>P. ovale</italic>. Its reported prevalence is generally low (<5%), except in West Africa, where prevalences above 10% have been observed in humans <xref ref-type="bibr" rid="pone.0005520-Cornu1">[6]</xref>, <xref ref-type="bibr" rid="pone.0005520-Faye1">[7]</xref>. <italic>P. ovale</italic> is often present in mixed infections and parasitaemia is usually low.</p><p><italic>P. ovale</italic> was previously thought to be present only in sub-Saharan Africa, Papua New Guinea, Irian Jaya in Indonesia and the Philippines <xref ref-type="bibr" rid="pone.0005520-Collins1">[4]</xref>. However, it appears to be more widely distributed, having been reported in the Middle East, the Indian Subcontinent and various parts of Southeast Asia <xref ref-type="bibr" rid="pone.0005520-Cadigan1">[8]</xref>–<xref ref-type="bibr" rid="pone.0005520-Incardona1">[11]</xref>. <italic>P. ovale</italic> has not been yet reported in South America. However, no global map of the geographical distribution of <italic>P. ovale</italic> has been produced since that of Lysenko and Beljaev in 1969 <xref ref-type="bibr" rid="pone.0005520-Lysenko1">[12]</xref>.</p><p>Few studies document the molecular diversity, geographical origin, evolutionary history and age of <italic>P. ovale</italic> populations. Based on complete DNA sequences of the small subunit ribosomal RNA (<italic>SSUrRNA</italic>) gene, partial sequences of cysteine protease, ookinete surface protein and <italic>cytochrome b</italic> genes, Win et al. (2004) compared <italic>P. ovale</italic> isolates from Myanmar, Indonesia and sequences available from GenBank. The result obtained supported the division of <italic>P. ovale</italic> into at least two types, but the classical and variant types identified did not differ morphologically and occurred in sympatry <xref ref-type="bibr" rid="pone.0005520-Win1">[13]</xref>, <xref ref-type="bibr" rid="pone.0005520-Tachibana1">[14]</xref>.</p><p>Phylogenetically, <italic>P. ovale</italic> clusters with <italic>Plasmodium</italic> species affecting simian primates (as do <italic>P. malariae</italic> and <italic>P. vivax</italic>, but not <italic>P. falciparum</italic>), but its phylogenetic relationships to other <italic>Plasmodium</italic> species or haemosporidian parasite genera remain unclear <xref ref-type="bibr" rid="pone.0005520-Collins1">[4]</xref>.</p><p>Three <italic>Plasmodium</italic> species, <italic>P. reichenowi</italic>, <italic>P. schwetzi</italic> and <italic>P. rodhaini</italic>, have already been reported in African great apes (chimpanzees and gorillas) and have been described as morphologically similar to <italic>P. falciparum</italic>, <italic>P. ovale</italic> or <italic>P. vivax</italic> (there are differing opinions) and <italic>P. malariae</italic>, respectively <xref ref-type="bibr" rid="pone.0005520-Coatney1">[15]</xref>. Like humans, the African great apes belong to the Hominidae family. Despite the close phylogenetic relationships between these non human primates and human hosts, the diversity of <italic>Plasmodium</italic> parasites in African great apes has been little studied and few molecular data for these parasites are available. Indeed, only one strain of <italic>P. reichenowi</italic>, originally isolated from a naturally infected chimpanzee (<italic>Pan troglodytes</italic>) in Central Africa (East of the Democratic Republic of the Congo) and adapted to a laboratory splenectomized chimpanzee, has been molecularly characterized <xref ref-type="bibr" rid="pone.0005520-Coatney1">[15]</xref>. This parasite is closely phylogenetically related to <italic>P. falciparum</italic>, the principal cause of human malaria. Data for other taxa, including genetically characterized non human primate malaria parasites, are required to provide insight into the evolutionary history of <italic>P. ovale</italic> <xref ref-type="bibr" rid="pone.0005520-Hagner1">[16]</xref>.</p><p>In order to investigate the diversity of <italic>Plasmodium</italic> parasites in African great apes, we screened 130 DNA samples from chimpanzees and gorillas in Cameroon. We found three chimpanzees infected by <italic>Plasmodium</italic> related to the human <italic>P. ovale</italic>. We present here the diversity of these chimpanzee parasites using two mitochondrial and one nuclear partial gene sequences and compared them to human <italic>P. ovale</italic> strains.</p></sec><sec id="s2"><title>Results</title><p>DNA samples from 130 chimpanzees and gorillas were tested for <italic>Plasmodium</italic> infection, using <italic>cytochrome b</italic> molecular tools. Two chimpanzees, CPZcam89 (225) and CPZcam91 (451), both belonging to subspecies <italic>Pan t</italic>. <italic>troglodytes</italic>, presented a single infection with <italic>Plasmodium</italic> parasites phylogenetically related to <italic>P. ovale</italic>. Both <italic>Plasmodium</italic> isolates were characterized by a unique DNA sequence for each of the <italic>cox1</italic>, <italic>cyt b</italic> and <italic>ldh</italic> markers, differing between the two isolates. A third chimpanzee (CPZcam63 (2360)), belonging to subspecies, <italic>Pan t</italic>. <italic>vellerosus</italic>, had a mixed infection composed of <italic>P. reichenowi</italic> and <italic>P. ovale</italic> related parasites. The latter has an identical <italic>cyt b</italic> sequence to <italic>Plasmodium</italic> found in CPZcam89 (451) chimpanzee; this isolate was discarded from the phylogenetic construction. The prevalence of <italic>P. ovale</italic> related <italic>Plasmodium</italic> species was found to be 2.3% (3/130) in the Cameroonian great apes tested. This prevalence is comparable to the prevalence of <italic>P. ovale</italic> in human populations from most endemic areas (<5%).</p><p>The 708 bp <italic>cyt b</italic> and the 964 bp <italic>cox1</italic> sequences as well as the 350 bp <italic>ldh</italic> sequence of the CPZcam89 (225) chimpanzee parasite strain are all identical to the human <italic>P. ovale</italic> variant type sequences (<xref ref-type="table" rid="pone-0005520-t001">Tables 1</xref>, <xref ref-type="table" rid="pone-0005520-t002">2</xref> and <xref ref-type="table" rid="pone-0005520-t003">3</xref>). Based on this genetic homology, this chimpanzee parasite strain was identified as being of the <italic>P. ovale</italic> variant type. The <italic>cyt b</italic>, <italic>cox1</italic> and <italic>ldh</italic> nucleotide sequences of the CPZcam91 (451) chimpanzee parasite diverged from the reported classical and variant <italic>P. ovale</italic> type nucleotide sequences (<xref ref-type="table" rid="pone-0005520-t001">Tables 1</xref>, <xref ref-type="table" rid="pone-0005520-t002">2</xref> and <xref ref-type="table" rid="pone-0005520-t003">3</xref>). For the <italic>cyt b</italic> marker, this chimpanzee <italic>Plasmodium</italic> sequence presented four synonymous mutations with respect to the classical <italic>P. ovale</italic> type sequence and one non synonymous mutation, M248I, with respect to the variant <italic>P. ovale</italic> type sequence (<xref ref-type="table" rid="pone-0005520-t001">Table 1</xref>). The <italic>cox1</italic> marker displayed two non synonymous mutations with respect to the classical <italic>P. ovale</italic> type and three with respect to the variant <italic>P. ovale</italic> type (<xref ref-type="table" rid="pone-0005520-t002">Table 2</xref>). The nuclear <italic>ldh</italic> sequence shows two non synonymous mutations compared to the classical <italic>P. ovale</italic> and four non synonymous mutations compared to the variant <italic>P. ovale</italic> (<xref ref-type="table" rid="pone-0005520-t003">Table 3</xref>).</p><table-wrap id="pone-0005520-t001" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0005520.t001</object-id><label>Table 1</label><caption><title>Substitutions and their positions in <italic>cyt b</italic> nucleotide sequences (numbers correspond to base pair positions and were defined according to the complete <italic>P. falciparum cyt b</italic> gene sequence M76611).</title></caption><graphic id="pone-0005520-t001-1" xlink:href="pone.0005520.t001"></graphic><table frame="hsides" rules="groups" alternate-form-of="pone-0005520-t001-1"><colgroup span="1"><col align="left" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col></colgroup><thead><tr><td align="left" rowspan="1" colspan="1">Sequences</td><td align="left" rowspan="1" colspan="1">315</td><td align="left" rowspan="1" colspan="1">375</td><td align="left" rowspan="1" colspan="1">402</td><td align="left" rowspan="1" colspan="1">450</td><td align="left" rowspan="1" colspan="1">492</td><td align="left" rowspan="1" colspan="1">510</td><td align="left" rowspan="1" colspan="1">514</td><td align="left" rowspan="1" colspan="1">534</td><td align="left" rowspan="1" colspan="1">744</td><td align="left" rowspan="1" colspan="1">756</td><td align="left" rowspan="1" colspan="1">774</td><td align="left" rowspan="1" colspan="1">885</td><td align="left" rowspan="1" colspan="1">903</td><td align="left" rowspan="1" colspan="1">948</td></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic> classical type</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic> variant type</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">A</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(M248I)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">CPZcam89 (225)</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">A</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(M248I)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">CPZcam91 (451)</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td></tr></tbody></table><table-wrap-foot><fn id="nt101"><p>Non synonymous mutation is shown in brackets.</p></fn></table-wrap-foot></table-wrap><table-wrap id="pone-0005520-t002" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0005520.t002</object-id><label>Table 2</label><caption><title>Substitutions and their positions in <italic>cox1</italic> nucleotide sequences (numbers correspond to base pair positions and were defined according to the complete sequence of the <italic>P. falciparum cox1</italic> gene M76611).</title></caption><graphic id="pone-0005520-t002-2" xlink:href="pone.0005520.t002"></graphic><table frame="hsides" rules="groups" alternate-form-of="pone-0005520-t002-2"><colgroup span="1"><col align="left" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col></colgroup><thead><tr><td align="left" rowspan="1" colspan="1">Sequences</td><td align="left" rowspan="1" colspan="1">449</td><td align="left" rowspan="1" colspan="1">458</td><td align="left" rowspan="1" colspan="1">462</td><td align="left" rowspan="1" colspan="1">473</td><td align="left" rowspan="1" colspan="1">575</td><td align="left" rowspan="1" colspan="1">632</td><td align="left" rowspan="1" colspan="1">650</td><td align="left" rowspan="1" colspan="1">657</td><td align="left" rowspan="1" colspan="1">761</td><td align="left" rowspan="1" colspan="1">764</td><td align="left" rowspan="1" colspan="1">765</td><td align="left" rowspan="1" colspan="1">766</td><td align="left" rowspan="1" colspan="1">830</td><td align="left" rowspan="1" colspan="1">966</td><td align="left" rowspan="1" colspan="1">1016</td><td align="left" rowspan="1" colspan="1">1022</td><td align="left" rowspan="1" colspan="1">1082</td></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic> classical type</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic> variant type</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">A</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(M211I)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">CPZcam89 (225)</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">A</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(M211I)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">CPZcam91 (451)</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">A</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(L255F)</td><td align="left" rowspan="1" colspan="1">(H256T)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr></tbody></table><table-wrap-foot><fn id="nt102"><p>Non synonymous mutations are shown in brackets.</p></fn></table-wrap-foot></table-wrap><table-wrap id="pone-0005520-t003" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0005520.t003</object-id><label>Table 3</label><caption><title>Substitutions and their positions in <italic>ldh</italic> nucleotide sequences (numbers correspond to base pair positions and were defined according to the complete sequence of the <italic>P. falciparum ldh</italic> gene PF13_0141).</title></caption><graphic id="pone-0005520-t003-3" xlink:href="pone.0005520.t003"></graphic><table frame="hsides" rules="groups" alternate-form-of="pone-0005520-t003-3"><colgroup span="1"><col align="left" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col></colgroup><thead><tr><td align="left" rowspan="1" colspan="1">Sequences</td><td align="left" rowspan="1" colspan="1">195</td><td align="left" rowspan="1" colspan="1">237</td><td align="left" rowspan="1" colspan="1">243</td><td align="left" rowspan="1" colspan="1">258</td><td align="left" rowspan="1" colspan="1">291</td><td align="left" rowspan="1" colspan="1">301</td><td align="left" rowspan="1" colspan="1">321</td><td align="left" rowspan="1" colspan="1">333</td><td align="left" rowspan="1" colspan="1">337</td><td align="left" rowspan="1" colspan="1">339</td></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic> classical type</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic> variant type</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">CPZcam89 (225)</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">CPZcam91 (451)</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">-</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(I113V)</td><td align="left" rowspan="1" colspan="1"></td></tr></tbody></table><table frame="hsides" rules="groups" alternate-form-of="pone-0005520-t003-3"><colgroup span="1"><col align="left" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col></colgroup><thead><tr><td align="left" rowspan="1" colspan="1">Sequences</td><td align="left" rowspan="1" colspan="1">351</td><td align="left" rowspan="1" colspan="1">361</td><td align="left" rowspan="1" colspan="1">387</td><td align="left" rowspan="1" colspan="1">406</td><td align="left" rowspan="1" colspan="1">427</td><td align="left" rowspan="1" colspan="1">475</td><td align="left" rowspan="1" colspan="1">504</td><td align="left" rowspan="1" colspan="1">507</td><td align="left" rowspan="1" colspan="1">510</td></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic> classical type</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic> variant type</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">C</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(S143P)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(K168N)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">CPZcam89 (225)</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">C</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(S143P)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(K168N)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">CPZcam91 (451)</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">T</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">-</td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">C</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">(F121L)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr></tbody></table><table-wrap-foot><fn id="nt103"><p>Non synonymous mutations are shown in brackets.</p></fn></table-wrap-foot></table-wrap><p>Investigation of the mitochondrial <italic>cyt b</italic>, <italic>cox1</italic> and nuclear <italic>ldh</italic> partial gene sequences in 13 <italic>P. ovale</italic> strains from humans from 12 different sites showed that <italic>P. ovale</italic> species could be divided into two distinct groups. Both classical and variant <italic>P. ovale</italic> (<xref ref-type="table" rid="pone-0005520-t004">Table 4</xref>) were associated with a unique sequence for each marker, consistent with the finding of Win et al, 2004 on <italic>cyt b</italic> gene <xref ref-type="bibr" rid="pone.0005520-Win1">[13]</xref>, <xref ref-type="bibr" rid="pone.0005520-Talman1">[17]</xref>. Comparisons of <italic>cyt b</italic> nucleotide sequences revealed 10 different substitutions between the variant and classical <italic>P. ovale</italic> types, one of which was a non synonymous mutation, M248I (<xref ref-type="table" rid="pone-0005520-t001">Table 1</xref>). Comparisons of the classical and variant <italic>cox1</italic> nucleotide sequences, also revealed 10 different mutations, one of which was a non synonymous mutation M211I (<xref ref-type="table" rid="pone-0005520-t002">Table 2</xref>). Comparisons of <italic>ldh</italic> classical and variant <italic>P. ovale</italic> nucleotide sequences showed 13 different substitutions, two of which were non synonymous mutations, S143P and K168N (<xref ref-type="table" rid="pone-0005520-t003">Table 3</xref>).</p><table-wrap id="pone-0005520-t004" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0005520.t004</object-id><label>Table 4</label><caption><title>Human <italic>P. ovale</italic> strains, strain code, geographical location of origin, nucleotide sequence, type and GenBank accession number.</title></caption><graphic id="pone-0005520-t004-4" xlink:href="pone.0005520.t004"></graphic><table frame="hsides" rules="groups" alternate-form-of="pone-0005520-t004-4"><colgroup span="1"><col align="left" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col></colgroup><thead><tr><td align="left" rowspan="1" colspan="1">Species</td><td align="left" rowspan="1" colspan="1">Strain code</td><td align="left" rowspan="1" colspan="1">Origin</td><td align="left" rowspan="1" colspan="1">GenBank accession number <italic>cytb</italic></td><td align="left" rowspan="1" colspan="1">GenBank accession number <italic>cox1</italic></td><td align="left" rowspan="1" colspan="1">Type</td></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">5894</td><td align="left" rowspan="1" colspan="1">Angola</td><td align="left" rowspan="1" colspan="1">FJ409567</td><td align="left" rowspan="1" colspan="1">FJ409571</td><td align="left" rowspan="1" colspan="1">classical</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">CAMBO</td><td align="left" rowspan="1" colspan="1">Cambodia</td><td align="left" rowspan="1" colspan="1">FJ409567</td><td align="left" rowspan="1" colspan="1">FJ409571</td><td align="left" rowspan="1" colspan="1">classical</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">3044</td><td align="left" rowspan="1" colspan="1">Republic of Central Africa</td><td align="left" rowspan="1" colspan="1">FJ409567</td><td align="left" rowspan="1" colspan="1">FJ409571</td><td align="left" rowspan="1" colspan="1">classical</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">5979</td><td align="left" rowspan="1" colspan="1">Ivory Coast</td><td align="left" rowspan="1" colspan="1">FJ409567</td><td align="left" rowspan="1" colspan="1">FJ409571</td><td align="left" rowspan="1" colspan="1">classical</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">3149</td><td align="left" rowspan="1" colspan="1">Gabon</td><td align="left" rowspan="1" colspan="1">FJ409567</td><td align="left" rowspan="1" colspan="1">FJ409571</td><td align="left" rowspan="1" colspan="1">classical</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">4646</td><td align="left" rowspan="1" colspan="1">Guinea</td><td align="left" rowspan="1" colspan="1">FJ409567</td><td align="left" rowspan="1" colspan="1">FJ409571</td><td align="left" rowspan="1" colspan="1">classical</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">3740</td><td align="left" rowspan="1" colspan="1">Democratic Republic of Congo</td><td align="left" rowspan="1" colspan="1">FJ409567</td><td align="left" rowspan="1" colspan="1">FJ409571</td><td align="left" rowspan="1" colspan="1">classical</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">4419</td><td align="left" rowspan="1" colspan="1">Cameroon</td><td align="left" rowspan="1" colspan="1">FJ409566</td><td align="left" rowspan="1" colspan="1">FJ409570</td><td align="left" rowspan="1" colspan="1">variant</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">5401</td><td align="left" rowspan="1" colspan="1">Madagascar</td><td align="left" rowspan="1" colspan="1">FJ409566</td><td align="left" rowspan="1" colspan="1">FJ409570</td><td align="left" rowspan="1" colspan="1">variant</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">2132</td><td align="left" rowspan="1" colspan="1">Mali</td><td align="left" rowspan="1" colspan="1">FJ409566</td><td align="left" rowspan="1" colspan="1">FJ409570</td><td align="left" rowspan="1" colspan="1">variant</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">5994</td><td align="left" rowspan="1" colspan="1">Mali</td><td align="left" rowspan="1" colspan="1">FJ409566</td><td align="left" rowspan="1" colspan="1">FJ409570</td><td align="left" rowspan="1" colspan="1">variant</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">2668</td><td align="left" rowspan="1" colspan="1">Rwanda</td><td align="left" rowspan="1" colspan="1">FJ409566</td><td align="left" rowspan="1" colspan="1">FJ409570</td><td align="left" rowspan="1" colspan="1">variant</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic></td><td align="left" rowspan="1" colspan="1">3043</td><td align="left" rowspan="1" colspan="1">Zimbabwe</td><td align="left" rowspan="1" colspan="1">FJ409566</td><td align="left" rowspan="1" colspan="1">FJ409570</td><td align="left" rowspan="1" colspan="1">variant</td></tr></tbody></table></table-wrap><p>The sequences presented are derived from a single PCR-sequencing event. The differences observed between these sequences, though likely to reflect reality, might be the result of PCR amplification artefacts.</p><p>Both of the methods used, maximum likelihood (ML) and Bayesian analyses, produced the same tree topology consistent with previous published <italic>Plasmodium</italic> phylogenetic analysis <xref ref-type="bibr" rid="pone.0005520-Escalante1">[18]</xref>, <xref ref-type="bibr" rid="pone.0005520-Hayakawa1">[19]</xref>. The phylogenetic relationships between the two <italic>Plasmodium</italic> strains isolated from chimpanzees to classical and variant <italic>P. ovale</italic> types, and the position of these strains within primate parasite group, are presented in <xref ref-type="fig" rid="pone-0005520-g001">Figure 1</xref>. The two chimpanzee parasites formed a monophyletic group with the two human <italic>P. ovale</italic> types. Monophyly was well supported by Bayesian posterior probabilities of 0.98 and a bootstrap value of 94%.</p><fig id="pone-0005520-g001" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0005520.g001</object-id><label>Figure 1</label><caption><title>Phylogeny of Haemosporidia inferred from <italic>cyt b</italic> and <italic>cox1</italic> nucleotide sequences.</title><p>Values are bootstrap percentages obtained by maximum likelihood analysis (left of the slash, values under 70% not shown) and Bayesian posterior probabilities (right of the slash, values less then 0.7 not shown), <italic>P</italic>. = <italic>Plasmodium</italic>. In red: Human malaria parasite species. Usual hosts are presented on the right side.</p></caption><graphic xlink:href="pone.0005520.g001"></graphic></fig></sec><sec id="s3"><title>Discussion</title><p>The characterization of 13 <italic>P. ovale</italic> human isolates, using mitochondrial <italic>cyt b</italic> and <italic>cox1</italic> markers and nuclear <italic>ldh</italic> marker from 12 different geographical locations, confirmed the diversification of human strains of <italic>P. ovale</italic> into two types, classical and variant <xref ref-type="bibr" rid="pone.0005520-Win1">[13]</xref>.</p><p>We reported here the first molecular finding of three chimpanzee <italic>Plasmodium</italic> isolates, one (CPZcam89 (225)) genetically identical to <italic>P. ovale</italic> variant type, one other (CPZcam91 (451)) closely related to human <italic>P. ovale</italic> types and a third one (CPZcam63 (2360)) showing mixed infection composed of <italic>P. reichenowi</italic> and <italic>P. ovale</italic> related parasite (the latter exhibits an <italic>cyt b</italic> sequence identical to CPZcam91 (451) <italic>cyt b</italic> sequence parasite). Phylogenetic analyses inferred from <italic>cyt b</italic> and <italic>cox1</italic> concatenates are well supported and show a monophyletic group composed of human <italic>P. ovale</italic> types and related chimpanzee parasites. The monophyly of the group is confirmed using <italic>ldh</italic> nuclear partial gene sequences (data not shown).</p><p><italic>P. schwetzi</italic> has been originally described by Reichenow in 1920 in blood apes in Cameroon <xref ref-type="bibr" rid="pone.0005520-Coatney1">[15]</xref>. <italic>P. schwetzi</italic> is morphologically similar to both <italic>P. vivax</italic> and <italic>P. ovale</italic> parasites that infect humans, and to date there are two equally convincing arguments to favour one or the other of these species as the most closely related to <italic>P. schwetzi</italic> <xref ref-type="bibr" rid="pone.0005520-Coatney1">[15]</xref>. Experimental infections by <italic>P. schwetzi</italic> in humans have also been reported <xref ref-type="bibr" rid="pone.0005520-Rodhain1">[20]</xref> and in 1970, Contacos established its potential as a zoonosis for Africa <xref ref-type="bibr" rid="pone.0005520-Contacos1">[21]</xref>. At present, no isolate of this parasite from which molecular sequences can be obtained is available.</p><p><italic>P. schwetzi</italic> often occurs as a mixed infection with <italic>P. reichenowi</italic> and <italic>P. rodhaini</italic>, the two other African great ape <italic>Plasmodium</italic> species described morphologically similar to <italic>P. falciparum</italic> and <italic>P. malariae</italic> respectively. In this study, we found one chimpanzee co-infected with <italic>P. reichenowi</italic> and a <italic>P. ovale</italic> related parasite molecularly identical to CPZcam91 (451) isolate. The CPZcam91 (451) chimpanzee parasite might be identified as being <italic>P. schwetzi</italic> regarding reports available on this species. Nevertheless, there is not enough evidence to support this. Morphological and other molecular information are needed to establish the identity of this parasite.</p><p>The identical sequences of CPZcam89 (225) chimpanzee parasite strain to the <italic>P. ovale</italic> variant type on both mitochondrial <italic>cyt b</italic> and <italic>cox1</italic> and nuclear <italic>ldh</italic> markers suggest possible cross-species transmission between human and chimpanzee hosts in Cameroon. Interestingly, a prevalence of <italic>P. ovale</italic> higher than that usually reported in Africa (above 10%) has been reported in two villages in the Manyemen forest province in Cameroon, where humans and great apes live in sympatry <xref ref-type="bibr" rid="pone.0005520-Cornu1">[6]</xref>. Furthermore, earlier, Lysenko and Beljaev (1969) previously reported a close relationship between <italic>P. ovale</italic> prevalence in humans and proximity to great apes in Africa <xref ref-type="bibr" rid="pone.0005520-Lysenko1">[12]</xref>.</p><p>No direct evidence for human malaria parasite transmission between apes and humans was reported in Gabon <xref ref-type="bibr" rid="pone.0005520-Ollomo1">[22]</xref>, but natural transmissions of human malaria parasites to non human primates have been reported in South America. <italic>P. falciparum</italic>, <italic>P. vivax</italic> and <italic>P. malariae</italic> transmissions to wild monkeys of the rainforest in French Guyana <xref ref-type="bibr" rid="pone.0005520-Fandeur1">[23]</xref> and to Brazilian wild monkeys <xref ref-type="bibr" rid="pone.0005520-deCastroDuarte1">[24]</xref> have also been documented. Experimental transmission of <italic>P. ovale</italic> to chimpanzees via sporozoite inoculation has been reported <xref ref-type="bibr" rid="pone.0005520-Bray1">[25]</xref>.</p><p>This study provides the first evidence of human <italic>P. ovale</italic> variant type in chimpanzees in Cameroon. A large molecular epidemiology study would be required to improve the documentation of potential natural bidirectional transmission between chimpanzee and human populations living in sympatry, making it possible to evaluate the potential role of African great apes as a reservoir for <italic>P. ovale</italic> in West Africa. The question raised by Haydon et al. (2002) concerning the possibility of human <italic>Plasmodium</italic> species being permanently maintained in chimpanzee populations, from which infection is transmitted to human, remains to be explored <xref ref-type="bibr" rid="pone.0005520-Haydon1">[26]</xref>.</p></sec><sec sec-type="materials|methods" id="s4"><title>Materials and Methods</title><sec id="s4a"><title>Chimpanzee and gorilla DNA specimens</title><p>Chimpanzees and gorillas, originated from different areas of Cameroon, were, for the most part, initially kept as pets for a variable period of time and then either brought to the local zoos or sanctuaries or confiscated by the Ministry of Environment and Forestry, then gathered in captivity. These animals were sampled and included during virological studies lead by the Virology Unit of Centre Pasteur du Cameroon <xref ref-type="bibr" rid="pone.0005520-Macfie1">[27]</xref>, <xref ref-type="bibr" rid="pone.0005520-Calattini1">[28]</xref>. A DNA bank was constituted between 1998 and 2004.</p><p>In total, we tested 130 DNA samples from great apes for <italic>Plasmodium</italic> infection, using <italic>cytochrome b</italic> (<italic>cyt b</italic>) molecular tools: 105 chimpanzees from 4 subspecies (60 <italic>Pan t</italic>. <italic>troglodytes</italic>, 39 <italic>Pan t</italic>. <italic>vellerosus</italic>, 3 <italic>Pan t</italic>. <italic>schweinfurthii</italic> and 3 <italic>Pan t</italic>. <italic>verus</italic>), 8 chimpanzees of undetermined subspecies and 17 gorillas (<italic>Gorilla gorilla</italic>).</p><p>Detailed information on the three positive samples: CPZcam89 (225): <italic>Pan t</italic>. <italic>troglodytes</italic> subspecies, juvenile female, collected in February 2000; CPZcam 91 (451): <italic>Pan t</italic>. <italic>troglodytes</italic> subspecies, adult male, collected in February 2001; CPZcam63 (2360): <italic>Pan t</italic>. <italic>vellerosus</italic> subspecies, adult male, collected in September 1998.</p></sec><sec id="s4b"><title><italic>Cyt b</italic> PCR amplification</title><p>We amplified 708 bp <italic>Cyt b</italic> gene fragments with two sets of primers, one for PCR reaction, PLAS1 (<named-content content-type="gene">5′-GAGAATTATGGAGTGGATGGTG-3′</named-content>) and PLAS2a (<named-content content-type="gene">5′-GTGGTAATTGACATCCWATCC-3′</named-content>) and one for nested-PCR, PLAS3 (<named-content content-type="gene">5′-GGTGTTTYAGATAYATGCAYGC-3′</named-content>) and PLAS4 (<named-content content-type="gene">5′-CATCCWATCCATARTAWAGCATAG-3′</named-content>) <xref ref-type="bibr" rid="pone.0005520-Duval1">[29]</xref>.</p><p>These primers are specifics for Haemosporidia parasites and do not amplify DNA from other Apicomplexa parasites or host DNA. PCR and nested-PCR were carried out in a final volume of 25 μl, under the following conditions: 2.5 μl of each primer (10 pmol/μl), 2 mM of each dNTP, 0.5 U of <italic>Taq</italic> polymerase (Solis), 2 mM MgCl<sub>2</sub> and 2 μl of DNA, heating for 5 minutes at 94°C, 30 s at 94°C, 30 s at 55°C and 1 min 30 s at 72°C for 40 cycles and a final extension phase for 10 minutes at 72°C. The PCR products were sequenced by Macrogen (Korea) using PLAS3 and PLAS4 primers.</p><p>The parasites isolated from African great apes were also characterized molecularly by another gene, the <italic>cytochrome c oxidase 1</italic> gene (<italic>cox1</italic>). This mitochondrial gene has been chosen for the international barcoding programme for biodiversity identification <xref ref-type="bibr" rid="pone.0005520-Hajibabaei1">[30]</xref>. Like <italic>cyt b</italic>, it is a conserved gene and is useful for resolving phylogenetic relationships between populations of parasite species that have diverged over tens or hundreds of millions of years <xref ref-type="bibr" rid="pone.0005520-Perkins1">[31]</xref>, <xref ref-type="bibr" rid="pone.0005520-Martinsen1">[32]</xref>.</p></sec><sec id="s4c"><title><italic>Cox1</italic> PCR amplification</title><p>We amplified 964 bp <italic>Cox1</italic> gene fragments with the PCR primer set, <italic>cox1a</italic>: <named-content content-type="gene">5′-CGCCTGACATGGATGGATAATAC -3′</named-content> and <italic>cox1b</italic>: <named-content content-type="gene">5′-CCATTTAAAGCGTCTGGATAATC -3′</named-content> and the nested-PCR primer set, <italic>cox1c</italic>: <named-content content-type="gene">5′-GATTAACCGCTGTCGCTGGGACTG -3′</named-content> and <italic>cox1d</italic>: <named-content content-type="gene">5′-CGTCTAGGCATTACATTAAATCC -3′</named-content>.</p><p>These primers are specifics of Haemosporidia parasites and do not amplify DNA from other Apicomplexa parasites or host DNA. PCR and nested-PCR were carried out in a final volume of 25 μl, under the following conditions: 2.5 μl of each primer (10 pmol/μl), 2 mM of each dNTP, 0.5 U of <italic>Taq</italic> polymerase (Solis), 1.5 mM MgCl<sub>2</sub> and 2 μl of DNA, 5 minutes at 94°C, 30 s at 94°C, 30 s at 53°C for PCR and 30 s at 58°C for nested-PCR, and 2 minutes at 72°C for 40 cycles, with a final extension period of 10 minutes at 72°C. The PCR products were sequenced by Macrogen (Korea) using <italic>cox1c</italic> and <italic>cox1d</italic> primers.</p><p>The nuclear lactate dehydrogenase (<italic>ldh</italic>) gene has also been used to characterize parasites isolated from chimpanzees.</p></sec><sec id="s4d"><title><italic>Ldh</italic> PCR amplification</title><p>We amplified 350 bp <italic>ldh</italic> gene fragments with two sets of primers, one for PCR reaction, LDH1 (<named-content content-type="gene">5′-GGNTCDGGHATGATHGGAGG-3′</named-content>) and LDH2 (<named-content content-type="gene">5′-GCCATTTCRATRATDGCAGC-3′</named-content>) and one for nested-PCR, LDH7 (<named-content content-type="gene">5′-TGTDATGGCWTAYTCVAATTGYMARGT-3′</named-content>) and LDH8 (<named-content content-type="gene">5′-CCATYTTRTTNCCATGWGCWSCDACA-3′</named-content>) <xref ref-type="bibr" rid="pone.0005520-Talman1">[17]</xref>.</p><p>These primers are specifics for Haemosporidia parasites and do not amplify DNA from other Apicomplexa parasites or host DNA. PCR and nested-PCR were carried out in a final volume of 25 μl, under the following conditions: 2.5 μl of each primer (10 pmol/μl), 2 mM of each dNTP, 0.5 U of <italic>Taq</italic> polymerase (Solis), 2,5 mM MgCl<sub>2</sub> and 2 μl of DNA ,heating for 5 minutes at 94°C, 30 s at 94°C, 30 s at 55°C for PCR and 30s at 52°C for nested-PCR, and 1min at 72°C for 40 cycles and a final extension phase for 10 minutes at 72°C. The PCR products were sequenced by Macrogen (Korea) using LDH7 and LDH8 primers.</p></sec><sec id="s4e"><title><italic>P. ovale</italic> human strains</title><p>We also characterized <italic>P. ovale</italic> from 12 isolates collected from 11 different African locations and 1 isolate collected from South-East Asia, Cambodia (<xref ref-type="table" rid="pone-0005520-t004">Table 4</xref>), in collaboration with the National Reference Center for Malaria (AP-HP, Hôpital Bichat-Claude Bernard, Paris, France) using the <italic>cyt b</italic>, <italic>cox1</italic> and <italic>ldh</italic> partial gene sequences.</p></sec><sec id="s4f"><title>Phylogenetic analyses</title><p>The <italic>cyt b</italic>, <italic>cox1</italic> and <italic>ldh</italic> sequences were checked using chromatograms and CLUSTALW alignment to ensure that none of the positions was ambiguous <xref ref-type="bibr" rid="pone.0005520-Thompson1">[33]</xref>. Mixed infection was discarded from the phylogenetic study. Phylogenetic analyses were based on the use of 708 bp <italic>cyt b</italic> and 964 bp <italic>cox1</italic> concatenated sequences (<xref ref-type="table" rid="pone-0005520-t005">Table 5</xref>). Reference sequences without ambiguous positions for either <italic>cyt b</italic> or <italic>cox1</italic> were retrieved from GenBank.</p><table-wrap id="pone-0005520-t005" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0005520.t005</object-id><label>Table 5</label><caption><title>Parasite taxa, with host name, geographical location and GenBank accession number of the <italic>cyt b</italic> and <italic>cox1</italic> sequences used for the phylogenetic analysis</title></caption><graphic id="pone-0005520-t005-5" xlink:href="pone.0005520.t005"></graphic><table frame="hsides" rules="groups" alternate-form-of="pone-0005520-t005-5"><colgroup span="1"><col align="left" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col></colgroup><thead><tr><td align="left" rowspan="1" colspan="1">Parasites</td><td align="left" rowspan="1" colspan="1">Host</td><td align="left" rowspan="1" colspan="1">Geographical location</td><td align="left" rowspan="1" colspan="1">GenBank accession number <italic>cyt b</italic></td><td align="left" rowspan="1" colspan="1">GenBank accession number <italic>cox1</italic></td></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>P. falciparum</italic></td><td align="left" rowspan="1" colspan="1"><italic>Homo sapiens</italic></td><td align="left" rowspan="1" colspan="1">Tropical regions</td><td align="left" rowspan="1" colspan="1">M76611</td><td align="left" rowspan="1" colspan="1">M76611</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. gonderi</italic></td><td align="left" rowspan="1" colspan="1">Old World monkeys</td><td align="left" rowspan="1" colspan="1">Central Africa</td><td align="left" rowspan="1" colspan="1">AY800111</td><td align="left" rowspan="1" colspan="1">AY800111</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. knowlesi</italic></td><td align="left" rowspan="1" colspan="1">Old World monkeys</td><td align="left" rowspan="1" colspan="1">Malaysia</td><td align="left" rowspan="1" colspan="1">AY598141</td><td align="left" rowspan="1" colspan="1">AY598141</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. malariae</italic></td><td align="left" rowspan="1" colspan="1"><italic>Homo sapiens</italic></td><td align="left" rowspan="1" colspan="1">Tropical and subtropical regions</td><td align="left" rowspan="1" colspan="1">AF069624</td><td align="left" rowspan="1" colspan="1">AF182848</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. vivax</italic></td><td align="left" rowspan="1" colspan="1"><italic>Homo sapiens</italic></td><td align="left" rowspan="1" colspan="1">Tropical and subtropical regions</td><td align="left" rowspan="1" colspan="1">AY598139</td><td align="left" rowspan="1" colspan="1">AY598139</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. simiovale</italic></td><td align="left" rowspan="1" colspan="1">Old World monkeys</td><td align="left" rowspan="1" colspan="1">Asia</td><td align="left" rowspan="1" colspan="1">AY800109</td><td align="left" rowspan="1" colspan="1">AY800109</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. simium</italic></td><td align="left" rowspan="1" colspan="1">New World monkeys</td><td align="left" rowspan="1" colspan="1">South America</td><td align="left" rowspan="1" colspan="1">AY800110</td><td align="left" rowspan="1" colspan="1">AY800110</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. cynomolgi</italic></td><td align="left" rowspan="1" colspan="1">Old World monkeys</td><td align="left" rowspan="1" colspan="1">Southeast Asia</td><td align="left" rowspan="1" colspan="1">AY800108</td><td align="left" rowspan="1" colspan="1">AY800108</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic> classical</td><td align="left" rowspan="1" colspan="1"><italic>Homo sapiens</italic></td><td align="left" rowspan="1" colspan="1">Tropical regions</td><td align="left" rowspan="1" colspan="1">FJ409567</td><td align="left" rowspan="1" colspan="1">FJ409571</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. ovale</italic> variant</td><td align="left" rowspan="1" colspan="1"><italic>Homo sapiens</italic></td><td align="left" rowspan="1" colspan="1">Tropical regions</td><td align="left" rowspan="1" colspan="1">FJ409566</td><td align="left" rowspan="1" colspan="1">FJ409570</td></tr><tr><td align="left" rowspan="1" colspan="1">CPZcam89 (225)</td><td align="left" rowspan="1" colspan="1"><italic>Pan t. troglodytes</italic></td><td align="left" rowspan="1" colspan="1">Tropical regions</td><td align="left" rowspan="1" colspan="1">FJ409565</td><td align="left" rowspan="1" colspan="1">FJ409569</td></tr><tr><td align="left" rowspan="1" colspan="1">CPZcam91 (451)</td><td align="left" rowspan="1" colspan="1"><italic>Pan t. troglodytes</italic></td><td align="left" rowspan="1" colspan="1">Tropical regions</td><td align="left" rowspan="1" colspan="1">FJ409564</td><td align="left" rowspan="1" colspan="1">FJ409568</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. yoelii</italic></td><td align="left" rowspan="1" colspan="1"><italic>Thamnomys rutilans</italic></td><td align="left" rowspan="1" colspan="1">Central Africa</td><td align="left" rowspan="1" colspan="1">M29000</td><td align="left" rowspan="1" colspan="1">M29000</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td align="left" rowspan="1" colspan="1"><italic>Grammomys surdaster</italic></td><td align="left" rowspan="1" colspan="1">Central Africa</td><td align="left" rowspan="1" colspan="1">AF014115</td><td align="left" rowspan="1" colspan="1">AF014115</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. chabaudi</italic></td><td align="left" rowspan="1" colspan="1"><italic>Thamnomys rutilans</italic></td><td align="left" rowspan="1" colspan="1">Central Africa</td><td align="left" rowspan="1" colspan="1">AF014116</td><td align="left" rowspan="1" colspan="1">AF014116</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. gallinaceum</italic></td><td align="left" rowspan="1" colspan="1"><italic>Gallus gallus</italic></td><td align="left" rowspan="1" colspan="1">Vietnam</td><td align="left" rowspan="1" colspan="1">AB250690</td><td align="left" rowspan="1" colspan="1">AB250690</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. relictum</italic></td><td align="left" rowspan="1" colspan="1">Birds</td><td align="left" rowspan="1" colspan="1">North America</td><td align="left" rowspan="1" colspan="1">AY099032</td><td align="left" rowspan="1" colspan="1">EU254593</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>P. juxtanucleare</italic></td><td align="left" rowspan="1" colspan="1"><italic>Gallus gallus</italic></td><td align="left" rowspan="1" colspan="1">Asia</td><td align="left" rowspan="1" colspan="1">AB250415</td><td align="left" rowspan="1" colspan="1">AB250415</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Leucocytozoon caulleryi</italic></td><td align="left" rowspan="1" colspan="1">Birds</td><td align="left" rowspan="1" colspan="1">Tropical regions</td><td align="left" rowspan="1" colspan="1">AB302215</td><td align="left" rowspan="1" colspan="1">AB302215</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Haemoproteus sp.</italic></td><td align="left" rowspan="1" colspan="1"><italic>Lichenostomus frenatus</italic></td><td align="left" rowspan="1" colspan="1">Australia</td><td align="left" rowspan="1" colspan="1">AY733087</td><td align="left" rowspan="1" colspan="1">AY733087</td></tr></tbody></table></table-wrap><p>Statistical analysis, based on the Xia and Xie method, was conducted to examine whether the number of substitutions was saturated or not <xref ref-type="bibr" rid="pone.0005520-Xia1">[34]</xref>. In this method, both transitions and transversions were plotted against evolutionary distances calculated with the JC69 model. The relative rates at which transitions and transversions saturated at the third position were compared by counting substitutions in all pairwise comparisons between sequences. The analysis showed that the third base was saturated, and this base was therefore discarded for subsequent phylogenetic analyses.</p><p>We identified the most appropriate nucleotide substitution model, based on hierarchical likelihood ratio tests (hLRTs), Akaike Information criterion (AIC) and bayesian information criterion (BIC) values, using PHYML <xref ref-type="bibr" rid="pone.0005520-Guindon1">[35]</xref> in a similar way to Modeltest <xref ref-type="bibr" rid="pone.0005520-Posada1">[36]</xref>. The Hasegawa, Kishino and Yano statistic HKY <xref ref-type="bibr" rid="pone.0005520-Hasegawa1">[37]</xref> was favoured by the hLRT and BIC tests. Rate variation between sites was allowed, with a gamma distribution for four rate categories for the nucleotide and amino acid data, allowing for invariant sites. Maximum likelihood and Bayesian trees were inferred using the previously described model. Maximum likelihood (ML) analysis was carried out with Phyml <xref ref-type="bibr" rid="pone.0005520-Guindon2">[38]</xref>, with nodal robustness evaluated by non-parametric bootstrapping (1000 replicates). Bayesian analysis was performed with MrBayes <xref ref-type="bibr" rid="pone.0005520-Huelsenbeck1">[39]</xref>, using two runs of 1 million generations sampled every 100 generations. Convergence was determined using the standard deviation of the split frequencies and runs were stopped when a value of less than 0.01 was reached. 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