Very little is known about the biting behavior of sylvan Anopheles and about the vertebrate hosts that constitute their preferred source of blood [ 33 ]. Zoophilic species, such as An. Our study suggests that these species also bite ungulates. This indicates that, concerning their blood meal, An.
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Such a propensity to bite a wide range of hosts is probably an adaptive trait in response to temporal fluctuations of host diversity and density in forest environments. This feature could enhance the possibility for cross-species transfer of parasites and could explain the parasite propensity to infect different host species.
In addition, the panel of Anopheles species infected by these newly described haemosporidian lineages encompassed mosquito species with a well-known anthropophilic feeding behavior sometimes very pronounced, such as in An. Finally phylogenetic analysis of the relationship between the Cyt-b sequences of the haemosporidian parasites obtained in our study and reference sequences S1 File indicated that the phylogenetic position of the four newly described haemosporidian lineages was closer to sauropsid Plasmodium and Polychromophilus bat parasites than to other mammalian parasites Fig 2 , Figure A in S1 File , and Figure B in S1 File.
According to the current Haemosporida classification, all species infecting mammals belong to different genera within the Plasmodiidae family [ 2 ]. However, molecular data are lacking for several genera identified in bats or flying squirrels e. Previous molecular-based studies proposed that most Plasmodiidae mammalian parasites i. It was suggested that parasites of the genus Polychromophilus in bats could be the result of a secondary invasion of mammals and that they derived from avian or reptile parasites sauropsid Plasmodium and were transmitted by bat flies [ 35 ].
The origin of Nycteria parasites is less clear and needs to be further investigated. Particularly, it is unclear whether they represent another case of host switch or an ancient mammalian Plasmodium lineage. Similarly, the origin of the four clades described in this study is uncertain. Indeed, depending on the phylogenetic analyses e. Therefore, it remains to be determined whether these four clades are the result of a host switch from sauropsid Plasmodium or whether they are more related to Polychromophilus parasites.
Another remaining question concerns the roles played by the vertebrate hosts and their vectors in their evolution and diversification. The bird lineage was related to parasites of the Haemoproteus genus [ 36 ] and the turtle lineage was related to parasites of the Haemocystidium genus [ 2 , 36 ]. More data on these groups of vertebrates are required to determine whether these last two lineages are new [ 2 , 25 , 36 ]. Indeed, the current data and those available in the literature are too scarce to conclude with confidence.
In this study, we identified four new haemosporidian molecular lineages that belong to the Plasmodiidae family and that might be transmitted by Anopheles mosquitoes. Concerning their vertebrate hosts, two of the lineages A and B can infect several mammalian species. The vertebrate hosts of the other two lineages C and D remain to be identified.
None of these four lineages could be matched against reference sequences, although several species of haemosporidian parasites were previously described in African antelopes, but only based on their morphological features. As a consequence, and in agreement with other authors e. Indeed, this genus includes some species that are more genetically distant between them than they are with species belonging to other genera. Ideally, this re-evaluation should be done using the different criteria of phenotypic similarities, the concept of biological species and, most importantly, the concept of phylogenetic species [ 2 , 10 ].
Such a re-evaluation is critical to our understanding of host-pathogen evolution in this group of parasites. The study was conducted in Gabon outside protected areas. All samples were collected from dead animals only bushmeat. Some animal carcasses confiscated from poachers by Gabonese authorities Fauna and Hunting Department, Ministry of Water Affairs and Forestry, Environment and Sustainable development of Gabon belonged to protected species e.
All samples obtained from bushmeat salesmen were collected only during the authorized hunting season of non-protected species and only in public markets. No money was given to the salesmen in exchange of the pieces of tissues from the animals. New samples were not collected for this study. At both sites Fig 1 , a longitudinal survey of sylvan Anopheles was carried out using CDC light traps placed in several sites of the forest between 5pm to 7am from October to December Mosquitoes were morphologically identified using identification keys [ 40 ].
Salivary glands of females Anopheles were separated from the rest of the body. In total, 2, females of Anopheles were collected that belonged to 17 species.
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Total DNA was extracted from bushmeat tissues, mosquito bodies and salivary glands with the DNeasy Blood and Tissue Kit Qiagen and used as template for the detection of haemosporidian parasites according to a previously described protocol [ 16 ] based on PCR amplification and sequencing of a portion of the parasite Cyt-b gene. The sequences reported in this study were deposited in GenBank under the following accession numbers KT to KT Because amplicons with chimeric sequences can form during PCR amplification of DNA from samples with multiple infections, similarity plot analyses were performed on the nucleotide alignments generated with the new and all reference sequences using the SIMPLOT package version 2.
Once verified that the amplicons did not contain chimeric sequences, phylogenetic analyses were done after multiple alignments of the obtained partial Cyt-b sequences nucleotides and of the Genbank reference sequences using ClustalW v 1. Software [ 42 ].
It was previously shown that enough phylogenetic data can be extracted from Cyt-b sequences to study the phylogenetic relationships between haemosporidian parasites and to recover major clades [ 43 , 44 ]. Maximum Likelihood ML methods were used for tree construction [ 16 ]. Recently, the choice of outgroup to use to root the tree of haemosporidian species has been a matter of debate. In our study, ML trees were rooted: i using Toxoplasma spp. Three independent runs were performed for 30 million generations sampled every 5, generations burn-in.
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One advantage of estimating a phylogeny using a relaxed clock is that an estimate of the position of the tree root can be obtained. The trees give the relationship between our sequences and others sequences of genbank. Phylogenetic tree was rooted using Leucocytozoon sp p Figure A.
Phylogenetic tree was rooted using Eimeria spp Figure B. And one table gives all accession numbers of sequences used in our study Table A. Authors thank the four reviewers for their comments on a previous version of the manuscript as well as Philippe Christe for helpful discussions. Browse Subject Areas? Click through the PLOS taxonomy to find articles in your field. Abstract Re-examination, using molecular tools, of the diversity of haemosporidian parasites among which the agents of human malaria are the best known has generally led to rearrangements of traditional classifications.
Introduction The order Haemosporida also called Haemosporidia, Haemospororida or Haemospororina includes many protozoan parasites among which the best known are the agents of human malaria, a disease affecting every year several millions of people in tropical regions and particularly in sub-Saharan Africa [ 1 ]. Results and Discussion In this study, we investigated the diversity of haemosporidian parasites that circulate among wild vertebrates in Gabon, Central Africa, using molecular tools to analyze the parasite content of bushmeat samples collected in different areas of Gabon Fig 1 and stored in a biobank.
Download: PPT. Fig 1. Fig 2. Phylogenetic relationships between the Cyt-b sequences of haemosporidian parasites obtained in our study and reference Cyt-b sequences names in black from existing databases. Table 1. Host species screened for the presence of haemosporidian parasites, number of tested vertebrate samples and number of specimens harboring a parasitic cytochrome B Cyt-b gene sequence and parasite lineage.
Fig 3. Conclusion In this study, we identified four new haemosporidian molecular lineages that belong to the Plasmodiidae family and that might be transmitted by Anopheles mosquitoes. Ethic statement. Molecular analyses Total DNA was extracted from bushmeat tissues, mosquito bodies and salivary glands with the DNeasy Blood and Tissue Kit Qiagen and used as template for the detection of haemosporidian parasites according to a previously described protocol [ 16 ] based on PCR amplification and sequencing of a portion of the parasite Cyt-b gene.
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S1 File. Phylogenetic trees and Accession numbers of all sequences. Acknowledgments Authors thank the four reviewers for their comments on a previous version of the manuscript as well as Philippe Christe for helpful discussions. References 1. Geneva World Health Organization Perkins SL.
Malaria's many mates: past, present, and future of the systematics of the order Haemosporida. The Journal of parasitology.
The origins of malaria: there are more things in heaven and earth. Further observations on the blood parasites of birds in Uganda. Journal of wildlife diseases. Garnham PC. Immunity against the different stages of malaria parasites. Bulletin de la Societe de pathologie exotique et de ses filiales. The chiropteran haemosporidian Polychromophilus melanipherus: a worldwide species complex restricted to the family Miniopteridae.
Infection, genetics and evolution: journal of molecular epidemiology and evolutionary genetics in infectious diseases. Study of gametocytes from the Plasmodium "vivax" group: morphology, development in Anopheles and infectivity of Plasmodium yoelii microgametocytes. Annales de parasitologie humaine et comparee. New data on the biology of gametocytes of Plasmodium yoelii yoelli gathered from morphological characteristics indicating their age.
Comptes rendus des seances de l'Academie des sciences Serie D, Sciences naturelles. Dionisia bunoi n. Species concepts and malaria parasites: detecting a cryptic species of Plasmodium. Manwell RD. Malaria infections by four species of plasmodium in the duck and chicken, and resulting parasite modifications.
American Journal of Epidemiology. View Article Google Scholar Jordan HB. The Journal of protozoology. Morphologically defined subgenera of Plasmodium from avian hosts: test of monophyly by phylogenetic analysis of two mitochondrial genes. Temporal dynamics and diversity of avian malaria parasites in a single host species. J Anim Ecol. Origin of the human malaria parasite Plasmodium falciparum in gorillas. African great apes are natural hosts of multiple related malaria species, including Plasmodium falciparum.
On the diversity of malaria parasites in African apes and the origin of Plasmodium falciparum from Bonobos. PLoS pathogens. The origin of malignant malaria. Plasmodium falciparum is not as lonely as previously considered. Ubiquitous Hepatocystis infections, but no evidence of Plasmodium falciparum-like malaria parasites in wild greater spot-nosed monkeys Cercopithecus nictitans. International journal for parasitology. By Ernest Folefack. Ernest Folefack holds an LL. Form of the State. Historical Note. Structure of Government. Applicable Law. National Legislation.
Regional Economic, Monetary and Business Law. Regional Economic and Monetary Law. Regional Business Law. Insurance Law. International Treaties Law. Legal Education and Legal Profession. Legal Education. Legal Profession. Legal Information. Selected Bibliography. Gabon is formally known as the Gabonese Republic. It won independence from the French on the 17 th august It is situated to the west of the Central Africa region. It is bordered to the north by Cameroon, to the east and south by the Republic of Congo, to the north-west by Equatorial Guinea, and to the west by the Atlantic Ocean.
It has a surface area of , sq km and a population of 1,, It is one of the least densely inhabited countries of Africa. It is endowed with a lot of mineral resources such as petroleum, uranium, phosphate, manganese, diamond, zinc, marble, niobium, and also very precious timber. It is inhabited mainly by about 40 Bantu ethnic groups, among which the Fang number around Form of the state. Gabon is a unitary state. It is divided into three levels of administrative units: 9 provinces, 47 divisions, and 26 sub-divisions or districts. The currency is the franc.
Before becoming a French colony, the Atlantic coast of this territory was constantly visited by the Portuguese since Since Dutch also took interest in the territory as slave trade was practiced with coastal local ethnic sovereigns, particularly the Mpongwe as intermediaries. Conflicts erupted from time with the Dutch on the control of the coastal islands and mainland but no exploration of the interior was carried out.
Exploration and penetration by the church effectively started in when the king of the Mpongwe signed a treaty for protection with the French. More French explorers went deeper into the land and signed treaties with tribal chiefs. In , the consolidated territories were made a French colony by decree. When France was occupied and politically divided in , French citizens living in Gabon fought a brief battle which ended in favour of the Gaullists and Gabon joined the allies. After the war, Gabonese started a struggle for independence. In , an internal autonomy was given to Gabon within the framework of the then French Community.
Independence was granted on the 17 th August He was an ex-service man of the French army and intelligence. He has since dominated the political life of Gabon even after the return to a multiparty system under pressure in The Constitution adopted in has been profoundly modified, and even reformed since It lays down a number of fundamental rights in favor of citizens but above all the organization and functioning of the three main branches of government.
There is a presidential system but it is formally called semi-presidential, with a President head of State elected by direct popular vote for a seven-year term no limits. The executive branch is the most powerful of the three branches. The President appoints a vice-President who is answerable to him alone and can not become president in case of vacancy.
The President also freely appoints a Prime minister who is theoretically head of Government as he proposes members of the Cabinet for appointment by the President. The legislative branch has a bicameral Parliament. The Senate consists of 91 seats elected for six-year terms by members of municipal councils and departmental or divisional assemblies. The National Assembly consists of seats. Members are elected for a five-year term by direct popular vote.
The legislative branch shares the right to initiate new laws with the executive. The judicial power is exercised by supreme courts, appeal courts, and lower courts.
The judiciary branch was profoundly modified and developed by the constitutional reform of Judicial Supreme Court Cour de Cassation. This is the Supreme Court which in French is referred to as ordinary affairs. It has jurisdiction on criminal, social, commercial, and civil matters.
It is divided into four chambers, each specializing in one of the areas of jurisdiction mentioned above. It receives cases in the last instance from appeal courts. This is the Supreme Court in administrative litigations or judicial review of the activities of the Executive and other public law bodies.
It also has advisory jurisdiction which allows the government to refer drafts of laws and regulations for legal advice before sending them to Parliament or to cabinet for signature. Accounting Supreme Court Cour des Comptes. This is the accounting court with jurisdiction to audit accounts of government and other public and semi-public bodies. It also has jurisdiction to adjudicate on all conflicts arising as the auditing of such bodies and to sanction the authorities concerned notwithstanding possible criminal investigations and prosecutions.
The Constitutional Court. Its new organization and jurisdiction was laid down by the constitutional reform, making it one of the most important Courts to support the rule of law. It is made up of nine judges appointed by: the President of the Republic 3 ; the President of the Senate 3 and the president of the National Assembly 3.
They have tenure of seven years, renewable once.