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The Primate Malarias

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The Primate Malarias 3 Ecology of the Hosts in Relation to the Transmission of Malaria ASIA BECAUSE the ecology of human malaria has been discussed by many Parts of Southeast Asia are the only areas authors, we will focus our main attention on the for which reasonably complete information is transmission biology of the malarias of non- available, and proven natural vectors of non- human primates; man and his malarias will be human primate malaria have been identified included, but only in those situations where the only from West Malaysia (Wharton and Eyles, two groups of primates and their parasites 1961; Wharton et al, 1962; Eyles et al, 1963; become part of the same transmission system. and Cheong et al, 1965). In 1960, the U.S. The level of speciation in the plasmodia of Public Health Service (NIH, LPC) established a non-human primates is high; 17 species have cooperative project with the Institute for been described from Asia, 3 from Africa (not Medical Research in Kuala Lumpur, Malaysia, including 2 from Madagascan lemurs), and 2 to investigate the potential for the natural from the New World. The fundamental ecology transmission of malaria parasites of monkeys to of the malarias of apes and monkeys is similar to man. The baseline information on the ecology of that of the human species. Susceptible malaria in the area was essentially complete mosquitoes must live for a sufficient period of because the personnel of the Institute for time to allow for completion of sporogony and Medical Research had maintained a high degree they must have a predilection for returning to the of interest in the bionomics of anophelines of the same group of vertebrates for a second blood Malay Peninsula for many years. meal. In addition, it is essential that the parasite The NIH-IMR team approached the be able to maintain an infection in the vertebrate problem on an ecological basis, studying host long enough for the vector to become separately, but by the use of essentially the same infected. Some degree of chronicity is required, techniques, the several distinct types of too, since either the rapid destruction of the environments found on the Malay Peninsula parasite by the vertebrate host or the rapid which supported monkeys and their malarias destruction of the vertebrate host by the parasite (Fig. 1). The basic techniques used in these would be detrimental, if not fatal, to the studies were described in detail by Wharton et al maintenance of the transmission system. Our (1963). Briefly, they consisted of monkey-baited approach to the discussion of the ecology of the net traps erected on platforms in the forest malarias of apes and monkeys has for canopy and run in conjunction with human- convenience and continuity been organized on a baited net traps, of similar design, on the ground geographical basis. nearby. Mosquito collections were made from these traps from sunset to sunrise, and all 19 20 PRIMATE MALARIAS FIGURE 1.—Vectors of human and animal malarias in various ecological zones in West Malaysia (modified from Wharton et al, 1964). anophelines taken in the traps were dissected nemestrina. The anopheline fauna in this area was and examined for malaria parasites. Sporozoite- dominated by members of the Anopheles umbrosus positive salivary glands were inoculated group. Natural malaria infections were known to intravenously into malaria-free rhesus monkeys. occur frequently in this species complex in The 3 basic types of environments in the area Malaysia and Hodgkin (1956) suspected that some are: fresh water swamp forests, primary rain of these infections might be of monkey origin. forests, and mangrove forests. As was expected, most of the anophelines Fresh Water Swamp Forests. This type caught in human- and monkey-baited net traps of forest covers a large portion of lowland belonged to the A. umbrosus group, however, A. Malaysia either as valleys, in which water letifer was the only species which invaded the collects during the rainy season, or extensive flat monkey-baited net traps in the canopy in significant areas, which maintain some level of water numbers. Sporozoites from a large number of through much of the year. These areas support naturally infected mosquitoes (including A. considerable populations of monkeys, including umbrosus and A. roperi as well as A. letifer) were both macaques (Macaca sp.) and langurs inoculated into malaria-free monkeys, but no (Presbytis sp.). Investigations were conducted in infections resulted. During the same period, a swamp forest north of Kuala Lumpur, experimental infections were being studied in the Malaysia where malaria infections had been laboratory, in Kuala Lumpur, to establish the level found in both Macaca fascicularis and M. of susceptibility of Malaysian anophelines to ECOLOGY OF THE HOSTS IN RELATION TO THE TRANSMISSION OF MALARIA 21 monkey malaria. Plasmodium cynomolgi (B A. leucosphyrus group of mosquitoes, are never far strain), which was originally isolated from M. from any location in West Malaysia. Finally, it irus (= fascicularis) from the east coast of should be noted that 20 A. pujutensis were trapped Malaysia near Kuantan (Garnham, 1959), was in the swamp forest environment but none was used as the laboratory parasite model for these found infected. This mosquito is of more than studies. Anopheles letifer from the swamp forest passing interest because it is a member of the A. study area, located on the opposite side of the leucosphyrus group in which 3 species are proven main Malaysian mountain range from the B vectors of simian malaria in Malaysia. strain type locality of P. cynomolgi, were Primary Rain Forests. The rain forests of exposed to experimental infections. Oocyst Southeast Asia occupy hill areas over parts of development was poor and no sporozoite Indochina, Thailand, Burma, Malaysia, Indonesia, differentiation was seen. These results seemed to and the Philippines, and all of these areas have preclude the involvement of A. letifer in the certain biological features in common. All support transmission of simian malaria even though this populations of non-human primates, and, wherever anopheline would invade the forest canopy to investigations have been carried out, non-human feed on monkeys. Eventually, the malaria primate malarias have been found. A typical high infections in the A. umbrosus group of canopied rain forest, in a mountainous area mosquitoes in this environment were attributed approximately 20 miles north and east of Kuala to P. traguli of the Malaysian mouse deer, Lumpur, was selected as a site for the investigation Tragulus javanicus, (Wharton et al, 1963). The of the transmission of simian malaria. A. letifer story was, however, not complete. Mosquito trapping activities in the canopy, When a strain of P. cynomolgi was isolated from with monkey bait, and on the ground, with human the swamp forest study area and exposed to A. bait, produced small catches both in number of letifer from the same locality, it was found that individuals and number of species represented. sporozoites developed readily in the salivary However, 2 species of anophelines, A. balabacensis glands. Such highly specific strain relationships introlatus and A. leucosphyrus, were attracted to between parasite and mosquito host must, on monkeys in the canopy and to man on the ground. occasion, be very important in the epidemiology Thus, 2 possible mosquito liaisons between the of monkey malaria. It is not known how malaria parasites of man and monkeys had been frequently such specific relationships occur, but found. It also seemed significant that both of these they could be an important consideration in any species belonged to the A. leucosphyrus group. area where simian malaria exists in the absence Members of this species complex were known to be of expected vectors or where expected vectors primarily jungle breeders, and had been seriously cannot be specifically incriminated in the incriminated as vectors of human malaria in parts of transmission. North Malaysia, Thailand, Cambodia, Indochina, The epidemiology of simian malaria in the Burma, and Borneo. Specimens of both species, fresh water swamp forest of Malaysia is with sporozoite positive salivary glands, were confused by the high levels of natural mosquito collected at the rain forest site. The sporozoites infections with P. traguli. It may be, that the from the naturally infected A. b. introlatus produced same species of anopheline is carrying both an infection of P. inui when inoculated simian and mouse deer malaria and it is intravenously into malaria-free rhesus monkeys impossible, so far, to find the former in a (Wharton et al, 1962). Using similar techniques, a mixture so dominated by the latter. At any rate, naturally infected A. leucosphyrus, trapped in the the vector of simian malaria in this environment same area, was found to harbor an infection with P. remains unknown. It is possible that the cynomolgi (Eyles et al, 1963). monkeys become infected in another area, The fact that 2 members of the same species because troops undergo a fairly constant complex were vectors of simian malaria was migratory activity, and forested hills, with particularly interesting when it was considered that potential breeding sites for proven vectors of the other species, in the same group, were believed to 22 PRIMATE MALARIAS transmit human malaria in areas where simian Investigations to determine the vector of the malaria was found, also. Studies to investigate monkey malaria were immediately successful. An the possibility that a single species might be A. hackeri, collected at the base of a nipah palm responsible for the transmission of both simian during a daytime-resting catch, was found to have and human malaria, in the same environment, sporozoite positive salivary glands. The sporozoites were undertaken in a forest area of Cambodia produced a P. knowlesi infection following where the local authorities had experienced inoculation into a malaria-free rhesus monkey serious difficulties in trying to prevent (Wharton and Eyles, 1961).
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