Simian malaria parasites of Southeast Asia: their Leucosphyrus Group natural vectors, hosts and geographical distribution (modified from Sallum et al 
Simian malaria parasites were first reported in Malayan monkeys by Daniels in 1908 . It had been assumed for a long time that transmission of simian malaria to humans would not be possible. However, an accidental infection of scientists in Atlanta, USA by mosquito bites in the laboratory proved that a simian malaria species–
Mosquito surveys carried out in the area where the first case occurred did not reveal any sporozoite infections in the mosquitoes. However, studies in the coastal areas of Selangor in peninsular Malaysia found
2. Simian malaria parasites and their hosts
In Southeast Asia, there are 13 species of
Information is currently available on the non-human primate malaria especially in Malaysia. Thus, so far five species of simian malaria parasites in non-human primates (macaques) have been reported from Malaysia [12, 14]. The simian malaria parasite
Simian malaria parasites have been detected in three main species of non-human primates. They are
The pig-tailed macaque –
3. History of natural infection of
P. knowlesi in human host
Scientists have always been curious as to the possibility of humans being infected with non-human primate malaria. This interest was intensified when two scientists working in the Memphis laboratory were infected with
In 1965, an American surveyor working in Bukit Kertau in Pahang, Malaysia came down with malaria. Fortunately he returned to USA where he was diagnosed as
4. Cases of knowlesi malaria in Southeast Asia
In 2004, a large focus of human knowlesi malaria cases were reported from Sarawak, Malaysian Borneo . In that study it was found that 58% of the patients, admitted at the Sarawak hospital, were found to be infected with knowlesi malaria using molecular tools. These were misidentified by microscopy as
After the publication in 2004 , more cases were reported in Malaysia [29-32] and also from other countries in Southeast Asia with the exception of Lao PDR. To date cases have been reported from Thailand [33-35], Philippines ,Vietnam , Indonesia , Cambodia , Myanmar  and Singapore . Malaysia has reported the highest number of cases in the region.
A study has shown that
Knowlesi malaria has shown to be life threatening and mortality has been reported [29, 31]. From December 2007 to November 2009 six (27%) out of 22 patients with severe knowlesi malaria died in Sabah . Cases of knowlesi malaria are also occurring in areas where human malaria cases have been reduced or in malaria free areas . People can contract malaria either outside their houses in rural settings, in farms where they work or in the forest while hunting or working.
5. Knowlesi malaria associated with travellers to Southeast Asia
Naturally acquired cases of
6. Bionomics of simian malaria vectors and trapping techniques
The distribution of
||Pf, Pv, Pm
|Indonesia, East Malaysia,
East Malaysia: Sarawak[45,75]
West Malaysia 
||Information inadequate||Luzon, Philippines|
||Probable vector of human malaria 
||Pf, Pv Pm
||India, Sri Lanka|
West Malaysia, Philippines, Thailand
||Probable vector of simian malaria parasites ||Indonesia, East and West Malaysia, Thailand|
As a member of the Leucosphyrus complex,
The Dirus complex is well known because its species are widespread in forest and forest foothills throughout the Oriental Region from southwestern India eastwards and from 30o north parallel to the Malaysian peninsula [60-62] (Figure2), whereas the Leucosphyrus complex has been investigated to a much lesser degree in Malaysia Borneo and Kalimantan Borneo.
6.2. Larval biology
Table 2 shows a summary of
||Heliophobic||Clear, turbid, fresh water
Muddy pool (W Malaysia )
|Still or stagnant||Small streams, seepage streams, pools||Wheel ruts, hoof prints|
||Typical heliophobic||Fresh water||Still or stagnant||Pools; dips in the ground||Wheel ruts, hoof prints|
||Heliophobic||Clear, turbid, fresh water||Still or stagnant||Small streams, pools, wells, dips in the ground||Borrow pits, wheel ruts, hoof prints|
||Heliophobic||Clean non saline water, but found in water containing up to 4% sea-water||Still or stagnant||In split bamboo and cavities at the leaf base of nipa palm||In Thailand, in elephant footprints |
6.3. Biological characteristics
The important biological charactersitics of the known vectors of simian malaria are shown in Table 3 which has been modified from Meek . Of the known vectors,
||Sarawak: Around midnight in forested areas and soon after dusk in village settlements 
Forest: 1900-2000 h; farm: 0100-0200 h 
Monkey biting rate at 6, 3 m above ground and at ground: 6.8:3.2:1.0. HBR highest at forest fringe (6.74%), within the forest (1.85%) and at long house (0.28%) 
|Similar host preference 1.0 : 1.3 ||Sarawak: parous rate 65.8% (farm), 53.7% (forest), 65.8% (longhouse) 
VC: 2.86 (farm), 0.60 (forest), 0.85 (longhouse) 
|Sarawak: 1.18% (pooled from forest fringe, forest & longhouse) , 0.7% (farm), 1.4% (forest), all confirmed Pk by PCR; EIR 11.98 (farm), 14.1 (forest) |
||Palawan: In and out; 20.00-03.00 h 
Sabah: 22:00-02:00 h [85- 86];after midnight 
Out (76%): 19:00-20:00 h, in(24%): 22:00-23:00 h;
Lombok: 19:00-21:00h) 
|Attracted to humans, monkey & water buffalo; more frequently caught in monkey traps ||Sabah: highest in Nov, lowest in July ||Kalimantan: 1.3% 
Palawan: 12.5% sporozoite rate; 29% attracted to monkeys were positive for oocysts 
||Late or early biting, usually around 22:00 h [60-62]
9 of 13
exophagic as well as endophagic and exophilic
|Higher parous rate (76%) & life expectancy during dry season compared to wet season (62.4%) in Lao ||Human sporozoite rates vary with season and location: from 7.8% in Assam (India) to 14% in Myanmar  and 2.5% in Laos 
43% of 72 salivary glands were PCR-positive for Pk CSP and Pk 18s rRNA. Mixed infections of Pk with Pv and Pf were common in Vietnam 
||Thailand: 1900-2100 h 
West Malaysia: 2000-2100 h; 74% biting before 2100 h; predominantly exophagic (1.11 bites/man-night) in both forest (1.24 bites/man-night) and fruit orchard (4.15 bites/man-night); 60% biting at ground level to 3 m high before 00:00 h; more biting at canopy level (6 m) compared to earlier collections at the same level .
|West Malaysia: 1: 2.6 [7.2]||West Malaysia: parous rate 65.7% (fruit orchard), 71.5% (forest)
VC: 2.46 (fruit orchard), 1.09 (forest) 
|West Malaysia: 0.60% (fruit orchard), 2.9% (forest)
EIR: 0.08 
||Not known since most bite monkeys and rarely found in human bait traps ||Most attracted to monkeys at canopy level in mangrove forest does not come to bite humans ||No data available||In coastal area of Rantau Panjang West Malaysia 0.7%
The peak biting times of
6.4. Laboratory susceptibility studies
In laboratory experiments with
6.5. Trapping techniques
Various trapping methods were tested for the collection of
This is similar to the human–bait-net trap introduced by Gater . This method provided the best results when tested . The platforms were constructed among the branches of trees to a height of 6 meters. Special metal cages measuring 90 cm x 90 cm x 90 cm and covered by wire mesh were used to house the monkeys on the platform measuring 300 cm X 200 cm. The meshed cages provided a physical barrier to prevent the monkeys from grabbing the collectors and also to prevent the entry of snakes. It is ideal to have two monkeys sharing a cage to increase vector attraction. A mosquito net measuring 190 cm x 180 cm x 150 cm with an opening of about 40 cm lifted on either ends was used to cover the cages with monkeys on each platform. The traps were operated from 18:00 to 06:00 hours and were searched at regular intervals [71, 72]. A collector, upon entering the net, closed the openings and collected all resting mosquitoes with the use of aspirators. Mosquitoes in the aspirator were then transferred to paper cups and were brought to the laboratory for identification and dissection. Platforms were built at various heights, ground level, 3 and 6 meters above ground. Figure 3 shows two different platforms in operation.
The other traps used were Shannon net trap, drum funnel-trap, Lumsden suction trap and light traps. Detailed descriptions can be found in Wharton . Of all the traps tested, it was found that the monkey- baited traps were superior compared to other types of traps. Although it is a difficult task to collect mosquitoes from the platforms at regular intervals, it is no doubt important to study the behaviour of the mosquitoes. Studies by Wharton  demonstrated that 83% of the
7. Implications for control
Currently insecticide treated bednets (ITN) and indoor residual spraying (IRS) are the two most important tools for the control of malaria vectors. Scaling up ITN, IRS, artemisinin-based combination therapies and intermittent preventive treatment for infants and pregnant women have contributed to the reported reductions in malaria on a global scale . As part of the Global Malaria Action Plan, the RBM Partnership and World Health Organization has recommended “malaria eradication worldwide by reducing the global incidence to zero through progressive malaria elimination in countries” . However, if human malaria could be eliminated, forests in Southeast Asia provide favourable environments for zoonotic transmission of
The vectors of
In Vietnam, forest malaria caused by
The use of repellents as personal protection measures have been advocated for malaria control. However, this needs to be evaluated in forest settings and large scale implementation will be a public health challenge. Among US Military troops, malaria cases have been reported due to non-compliance of personal protective measures and failure of chemoprophylaxis . Currently in Malaysia people are getting infected when they visit plantations or forests for work or recreational activities as some important vectors do not enter houses .
There is no reason to doubt the possibility and biological capacity of other simian malaria species to infect humans [13, 107].
Currently only three species of mosquitoes have been incriminated as simian malaria vectors in Malaysia (
In Thailand, the main vectors for human malaria are
According to Obsomer et al  the mean temperature below 20o C seems to limit the northern distribution of the Dirus complex to just beyond the border of India with Nepal and Bhutan. Rainfall is probably the limiting factor to the west with annual rainfall per year under 800 mm. Thus the lack of information on the distribution and occurrence of
Thus it is timely to determine all the vectors of simian malaria throughout the Southeast Asian region. Although old records stating the distribution of the various
Since many malaria control programmes in Southeast Asia are moving towards elimination of malaria , it is important to determine the prevalence of knowlesi malaria in these countries. In the Greater Mekong Subregion including Bangladesh and India
The authors thank Pollie Rueda for his constructive comments. The first author was supported by a grant from University of Malaya UM.C/625/1/HIR/099.J-20011-73822
Daniels CW. Animal parasites in man and some of the lower animals in Malaya. Studies Institite for Medical Research FMS 1908;3:1-13.
Coatney GR. Simian malaria: Its importance to world wide eradication of malaria. Journal of American Medical Association 1963;184:876-7.
Contacos PG, Elder HA, Coatney GR. Man to man transfer of two strains of Plasmodium cynomolgiby mosquito bites. American Journal of Tropical Medicine and Hygiene 1962;11:186-93.
Chin W, Contacos PG, Coatney GR, Kimball HR. A naturally acquired quotidian-type malaria in man transferable to monkeys. Science 1965;149:865.
Sinton JA MH. A critical review of the literature relating to the identification of the malaria parasites recorded from monkeys of the families Cercopithecidae and Colobidae. Rec Malaria Surv India 1932-1933;III:357–443.
Warren MW, Cheong WH, Fredericks HK, Coatney GR. Cycles of jungle malaria in West Malaysia. American Journal of Tropical Medicine and Hygiene 1970;19:383.
Fong YL, Cadigan FC, Coatney GR. A presumptive case of naturally occurring Plasmodium knowlesimalaria in man in Malaysia. Transactions of Royal Society of Tropical Medicine and Hygiene 1971;65:839.
Wharton RH, Eyles DE. Anopheles hackeri, a vector of Plasmodium knowlesiin Malaya. Science 1961;134:279.
Chin W, Contacos PG, Collins WE, Jeter MH, Alpert E. Experimental mosquito-transmission of Plasmodium knowlesito man and monkey. American Journal of Tropical Medicine and Hygiene 1968;17:355-8.
Singh B, Sung LK, Matusop A, Radhakrishnan A, Shamsul SSG, Cox-Singh J, et al. A large focus of naturally acquired Plasmodium knowlesiinfections in human beings. Lancet 2004;363:1017-24.
Collins WE. Major animal models in malaria research: Simian. In: Wernsdofer WH and McGregor editors. Malaria: Principles and Practice of Malariology. Edinburgh: Churchill Livingstone; 1988. p. 1473-501.
Fodden J. Malaria in Macaques. International Journal of Primatology 1994;15:573-96.
Galinski MR, Barnwell JW. Monkey malaria kills four humans. Trends in Parasitology 2009;25:200-4.
Lee KS, Divis PCS, Zakaria SK, Matusop A, Julin RA, Conway DJ, et al. Plasmodium knowlesi:reservoir hosts and tracking the emergence in humans and macaques. PLoS Pathogens 2011;7:e1002015.
Coatney GR, Elder HA, Contacos PG, Getz ME, Greenland R, Rossan RN, et al. Transmission of the M strain of Plasmodium cynomolgito man. The American Journal of Tropical Medicine and Hygiene 1961;10:673-8.
Collins WE, Warren MW, Sullivan JAS, Galland GG, Nace D, Williams A, et al. Studies on two strains of Plasmodium cynomolgiin New World and Old World monkeys and mosquitoes. Journal of Parasitology 2005;91:280-3.
Eyles DE, Laing ABG, Fong YL. Plasmodium fieldisp nov, a new species of malaria parasite from the pig-tailed macaque in Malaya. Annals of Tropical Medicine and Parasitology 1962;56:242-7.
Eyles DE. The species of simian malaria: Taxonomy, morphology, life cycle, and geographical distribution of the monkey species. Journal of Parasitology 1963:866-87.
Eyles DE, Laing ABG, Dobrovolny CG. The malaria parasites of the pig-tailed macaque, Macaca nemestrina(Linnaeus). Indian Journal of Malariology 1962;16:285-98.
Eyles DE, Laing ABG, Warren M, Sandosham AA. Malaria parasites of the Malayan leaf monkeys of the genus Presbytis. Medical Journal of Malay 1962;17:85-6.
Li M. Identification and molecular characterization of simian malaria parasites in wild monkeys of Singapore. Singapore: National University of Singapore; 2011.
Ho G, Lee C, Abie M, Zainuddin Z, Japnin J, Topani R, et al. Prevalance of Plasmodiumin the Long-tailored Macaque ( Macaca fasicularis) from Selangor, Malaysia. Proceedings 13th Association of Institutions for Tropical Veterinary Medicine (AITVM) Conference 2010:49.
Eyles DE, Fong, Y.l.,Warren, Mc,W., Guinn,E.,Sabdosham,AA.,Wharton,R.H. Plasmodium coatneyi,a new species of primate malaria from Malaya. American Journal of Tropical Medicine and Hygiene 1962;11:597-604.
Eyles DE, Dunn F, Warren McW, Guinn E. Plasmodium coatneyifrom the Philippines. Journal of Parasitology 1963;49:1038.
Sandosham A. Recent researchers on malaria at the Institute for Medical Research, Kuala Lumpur. Medical Journal of Malaysia 1967;22:145-60.
Lee KS, Cox-Singh J, Singh B. Morphological features and differential counts of Plasmodium knowlesiparasites in naturally acquired human infections. Malaria Journal 2009;8:73.
Coatney RG, Collins WE, Warren M, Contacos PG. The Primate Malarias. Washington: U.S. Government Printing Office; 1971.
Sandosham AA. Malariology with special reference to Malaya Singapore. Kuala Lumpur: University of Malaya Press; 1959.
Cox Singh J, Davis TME, Lee KS, Shamsul SSG, Matusop A, Ratnam S, et al. Plasmodium knowlesimalaria in humans is widely distributed and potentially life threatening. Clinical Infectious Diseases 2008;46:165-71.
Vythilingam I, NoorAzian YM, Huat TC, Jiram AI, Yusri YM, Azahari AH, et al. Plasmodium knowlesiin humans, macaques and mosquitoes in peninsular Malaysia. Parasite Vectors 2008;1:26.
William T, Menon J, Rajahram G, Chan L, Ma G, Donaldson S, et al. Severe Plasmodium knowlesimalaria in a tertiary care hospital, Sabah, Malaysia. Emerging Infectious Diseases 2011;17:1248-54.
Barber BE, William T, Jikal M, Jilip J, Dhararaj P, Menon J, et al. Plasmodium knowlesimalaria in children. Emerging Infectious Diseases 2011;17:20.
Jongwutiwes S, Putaporntip C, Iwasaki T, Sata T, Kanbara H. Naturally acquired Plasmodium knowlesimalaria in human, Thailand. Emerging Infectious Diseases 2004;10:2211-3.
Jongwutiwes S, Buppan P, Kosuvin R, Seethamchai S, Pattanawong U, Sirichaisinthop J, et al. Plasmodium knowlesiMalaria in humans and macaques, Thailand. Emerging Infectious Diseases 2011;17:1799-806.
Putaporntip C, Hongsrimuang T, Seethamchai S, Kobasa T, Limkittikul K, Cui L, et al. Differential prevalence of Plasmodiuminfections and cryptic Plasmodium knowlesimalaria in humans in Thailand. Journal of Infectious Diseases 2009;199:1143.
Luchavez J, Espino F, Curameng P, Espina R, Bell D, Chiodini P, et al. Human infections with Plasmodium knowlesi, the Philippines. Emerging Infectious Diseases 2008;14:811-3.
Eede P, Van H, Van Overmeir C, Vythilingam I, Duc T, Hung L, et al. Human Plasmodium knowlesiinfections in young children in central Vietnam. Malaria Journal 2009;8:249.
Sulistyaningsih E, Fitri LE, Löscher T, Berens-Riha N. Diagnostic difficulties with Plasmodium knowlesiinfection in humans. Emerging Infectious Diseases 2010;16:1033.
Khim N, Siv S, Kim S, Mueller T, Fleischmann E, Singh B, et al. Plasmodium knowlesiinfection in humans, Cambodia, 2007-2010. Emerging Infectious Diseases 2011;17:1900.
Zhu H, Li J, Zheng H. Human natural infection of Plasmodium knowlesi. Zhongguo ji sheng chong xue yu ji sheng chong bing za zhi= Chinese Journal Parasitology Parasite Diseases 2006;24:70.
Ng OT, Ooi EE, Lee CC, Lee PJ, Ng LC, Pei SW, et al. Naturally acquired human Plasmodium knowlesiinfection, Singapore. Emerging Infectious Diseases 2008;14:814.
White N. Plasmodium knowlesi: the fifth human malaria parasite. Clinical Infectious Diseases 2008;46:172.
Anderios F, NoorRain A, Vythilingam I. In vivo study of human Plasmodium knowlesiin Macaca fascicularis. Experimental Parasitology 2010;124:181-9.
Garnham P. Malaria parasites and other haemosporidia. Oxford: Blackwell Scientific Publications; 1966.
Vythilingam I. Review Paper Plasmodium knowlesiin humans: a review on the role of its vectors in Malaysia. Tropical Biomedicine 2010;27:1-12.
Hoosen A, Shaw M. Plasmodium knowlesiin a traveller returning to New Zealand. Travel Medicine and Infectious Disease 2011;9:144-8.
Mali S, Steele S, Slutsker L, Arguin PM. Malaria surveillance–United states, 2006. MMWR Surveill Summ 2008;57:24-39.
Kantele A, Marti H, Felger I, Muller D, Jokiranta TS. Monkey Malaria in a European Traveler Returning from Malaysia. Emerging Infectious Diseases 2008;14:1434-6.
Bronner U, Divis PCS, Farnert A, Singh B. Swedish traveller with Plasmodium knowlesimalaria after visiting Malaysian Borneo: a case report. Malaria Journal 2009;8:15.
Tang TH, Salas A, Ali-Tammam M, Martínez M, Lanza M, Arroyo E, et al. First case of detection of Plasmodium knowlesiin Spain by Real Time PCR in a traveller from Southeast Asia. Malaria Journal 2010;9:219.
Berry A, Iriart X, Wilhelm N, Valentin A, Cassaing S, Witkowski B, et al. Imported Plasmodium knowlesimalaria in a French tourist returning from Thailand. American Journal of Tropical Medicine and Hygiene 2011;84:535-8.
Warren MW, Wharton RH. The vectors of simian malaria: identity, biology, and geographical distribution. Journal of Parasitology 1963:892-904.
Sallum MAM, Peyton EL, Wilkerson RC. Six new species of the Anopheles leucosphyrusgroup, reinterpretation of An. elegansand vector implications. Medical and Veterinary Entomology 2005;19:158-99.
Harbach RE. The classification of genus Anopheles(Diptera: Culicidae): a working hypothesis of phylogenetic relationships. Bulletin of Entomological Research 2004;94:537-54.
Peyton E. A new classification for the Leucosphyrus Group of Anopheles( Cellia). Mosquito Systematics 1989;21:197-205.
Sallum MAM, Peyton EL, Harrison BA, Wilkerson RC. Revision of the Leucosphyrus group of Anopheles( Cellia)(Diptera, Culicidae). Revista Brasileira de Entomolo 2005;49:01-152.
Manguin S, Garros C, Dusfour I, Harbach RE, Coosemans M. Bionomics, taxonomy, and distribution of the major malaria vector taxa of Anophelessubgenus Celliain Southeast Asia: An updated review. Infection, Genetics and Evolution 2008;8:489-503.
Rattanarithikul R, Harrison BA. An illustrated key to the Anopheleslarvae of Thailand. United State Army Medical Component, South East Asia Treaty Organization, Bangkok 1973.
Rattanarithikul R, Harrison BA, Harbach RE, Panthusiri P, Coleman RE. Illustrated keys to the mosquitoes of Thailand. IV. Anopheles. Southeast Asian Journal of Tropical Medicine Public Health 2006;37 1-128.
Baimai V, Kijchalao U, Sawadwongporn P, Green CA. Geographic distribution and biting behaviour of four species of the Anopheles diruscomplex (Diptera: Culicidae) in Thailand. Southeast Asian Journal of Tropical Medicine Public Health 1988;19:151-61.
Obsomer V, Defourny P, Coosemans M. The Anopheles diruscomplex: spatial distribution and environmental drivers. Malaria Journal 2007;6:26.
Trung HD, Bortel WV, Sochantha T, Keokenchanh K, Briët OJT, Coosemans M. Behavioural heterogeneity of Anophelesspecies in ecologically different localities in Southeast Asia: a challenge for vector control. Tropical Medicine & International Health 2005;10:251-62.
Vythilingam I. Plasmodium knowlesiand Wuchereria bancrofti: their vectors and challenges for the future. Frontiers in Systems Biology 2012;3.
Sinka ME, Bangs MJ, Manguin S, Chareonviriyaphap T, Patil AP, Temperley WH, et al. The dominant Anophelesvectors of human malaria in the Asia-Pacific region: occurrence data, distribution maps and bionomic précis. Parasite Vectors 2011;4:1-46.
Reid JA. Anopheline Mosquitoes of Malaya and Borneo. Malaysia: Institute for Medical Research Malaysia; 1968.
Colless D. The Anopheles leucosphyrusgroup. Transactions of Royal Entomological Society London 1956;108:37-116.
Meek S. Vector control in some countries of Southeast Asia: comparing the vectors and the strategies. Annals of Tropical Medicine and Parasitology 1995;89:135-47.
Chang MS, Doraisingam P, Hardin S, Nagum N. Malaria and filariasis transmission in a village/forest setting in Baram District, Sarawak, Malaysia. Journal of Tropical Medicine and Hygiene 1995;98:192.
Colless DH. Observations on Anopheline mosquitoes of the Akah river, 4th Division, Sarawak. Bulletin of Entomological Research 1956;47:115-23.
Zulueta J. Malaria in Sarawak. Bulletin World Health Organisation 1956;15:651-71.
Tan CH, Vythilingam I, Matusop A, Chan ST, Singh B. Bionomics of Anopheles latensin Kapit, Sarawak, Malaysian Borneo in relation to the transmission of zoonotic simian malaria parasite Plasmodium knowlesi. Malaria Journal 2008;7:52.
Jiram AI, Vythilingam I, NoorAzian YM, Yusof YM, Azahari AH, Fong MY. Entomologic investigation of Plasmodium knowlesivectors in Kuala Lipis, Pahang, Malaysia. Malaria Journal 2012;11:213.
Tsukamoto M, Miyata A, Miyagi I. Surveys on simian malaria parasites and their vector in Palawan Island, the Philippines. Tropical Medicine 1978;20:39-50.
Miyagi I. Studies on malaria vectors in Philippines especially on Anopheles balabacensis balabacensisand monkey malaria in Palawan. The Journal of Tropical Medicine 1973;2:163.
Vythilingam I, Tan CH, Asmad M, Chan ST, Lee KS, Singh B. Natural transmission of Plasmodium knowlesito humans by Anopheles latensin Sarawak, Malaysia. Transactions Royal Society Tropical Medicine Hygiene 2006;100:1087-8.
Nakazawa S, Marchand RP, Quang NT, Culleton R, Manh ND, Maeno Y. Anopheles dirusco-infection with human and monkey malaria parasites in Vietnam. International Journal of Parasitology 2009;39:1533-7.
Marchand RP, Culleton R, Maeno Y, Quang NT, Nakazawa S. Co-infections of Plasmodium knowlesi, P. falciparum, and P. vivaxamong humans and Anopheles dirusmosquitoes, southern Vietnam. Emerging Infectious Diseases 2011;17:1232-9.
Wharton RH, Eyles DE, Warren M, Cheong WH. Studies to determine the vectors of monkey malaria in Malaya. Annals of Tropical Medicine and Parasitology 1964;58:56.
Tan CH. Identification of vectors of Plasmodium knowlesiand other malaria parasites, and studies on their bionomics in Kapit, Sarawak, Malaysia. MSc Thesis University Malaysia, Sarawak (UNIMAS) 2008.
Cheong WH, Warren MW, Omar AH, Mahadevan S. A nopheles balabacensis balabacensisidentified as vector of simian malaria in Malaysia. Science 1965;150:1314.
Vythilingam I, Phetsouvanh R, Keokenchanh K, Yengmala V, Vanisaveth V, Phompida S, et al. The prevalence of Anopheles(Diptera: Culicidae) mosquitoes in Sekong Province, Lao PDR in relation to malaria transmission. Tropical Medicine International Health 2003;8:525-35.
Pothikasikorn J, Bangs MJ, Boonplueang R, Chareonviriyaphap T. Susceptibility of various mosquitoes of Thailand to nocturnal subperiodic Wuchereria bancrofti. Journal of Vector Ecology 2008;33:313-20.
Vythilingam I, Chan ST, Shanmugratnam C, Tanrang H, Chooi KH. The impact of development and malaria control activities on its vectors in the Kinabatangan area of Sabah, East Malaysia. Acta Tropica 2005;96:24-30.
Maekawa Y, Tsuda Y, Dachlan YP, Yotopranoto S, Gerudug IK, Yoshinaga K, et al. Anopheline fauna and incriminatory malaria vectors in malaria endemic areas on Lombok Island, Indonesia. Medical Entomology and Zoology 2009;60:1-11.
Hii J. Evidence for the existence of genetic variability in the tendency of Anopheles balabacensisto rest in houses and to bite man. The Southeast Asian Journal of Tropical Medicine and Public Health 1985;16:173.
Rohani A, Lokman Hakim S, Hassan AR, Chan ST, Ong YF, Abdullah AG, Lee HL Bionomics of Anopheles balabacensisBaisas, the Principal Malaria Vector in Ranau, Sabah. Tropical Biomedicine 1999;16:31-8.
Chiang G, Cheong W, Samarawickrema W, Mak J, Kan S. Filariasis in Bengkoka Peninsula, Sabah, Malaysia: vector studies in relation to the transmission of filariasis. The Southeast Asian Journal of Tropical Medicine and Public Health 1984;15:179.
Schultz G. Biting activity of mosquitoes (Diptera: Culicidae) at a malarious site in Palawan, Republic of The Philippines. Southeast Asian Journal Tropical Medicine and Public Health 1992;23:464-9.
Harbach RE, Baimai V, Sukowati S. Some observations on sympatric populations of the malaria vectors Anopheles leucoshpyrusand Anopheles balabacensisin a village forest setting in South Kalimantan. Southeast Asian Journal of Tropical Medicine and Public Health 1987;18:241-7.
Sungvornyothin S, Kongmee M, Muenvorn V, Polsomboon S, Bangs MJ, Prabaripai A, et al. Seasonal abundance and bloodfeeding activity of Anopheles dirussensu lato in western Thailand. Journal of the American Mosquito Control Association 2009;25:425-30.
Van Bortel W, Trung HD, Hoi le X, Van Ham N, Van Chut N, Luu ND, Roelants P, et al. Malaria transmission and vector behaviour in a forested malaria focus in central Vietnam and the implications for vector control. Malaria Journal 2010;9:373.
Lambrechts L, Knox TB, Wong J, Liebman KA, Albright RG, Stoddard ST. Shifting priorities in vector biology to improve control of vector borne disease. Tropical Medicine & International Health 2009;14:1505-14.
Collins WE, Contacos PG, Guinn EG. Studies on the transmission of simian malarias II. Transmission of the H strain of Plasmodium knowlesiby Anopheles balabacensis balabacensis. Journal of Parasitology 1967:841-4.
Collins WE, Contacos PG, Guinn EG, Held JR. Studies on the transmission of simian malarias, I. Transmission of two strains of Plasmodium inuiby Anopheles maculatusand A. stephensi. Journal of Parasitology 1966:664-8.
Wharton RH, Eyles DE, Warren McW. The development of methods for trapping the vectors of monkey malaria. Annals of Tropical Medicine and Parasitology 1963;57:32-46.
Gater BAR. Aids to the identification of anopheline imagines in Malaya. Singapore: Government Printer; 1935.
Takken W, Knols BGJ. Malaria vector control: current and future strategies. Trends in Parasitology 2009;25:101-4.
WHO. The global malaria action plan for a malaria free world. http://wwwrollbackmalariaorg/gmap accessed 18Sept 2012. Geneva 2008.
Cox-Singh J, Singh B. Knowlesi malaria: newly emergent and of public health importance? Trends in Parasitology 2008;24:406-10.
Rajahram GS, Barber BE, William T, Menon J, Anstey NM, Yeo TW. Deaths due to Plasmodium knowlesimalaria in Sabah, Malaysia: association with reporting as Plasmodium malariaeand delayed parenteral artesunate. Malaria Journal 2012;11:284.
Ferguson HM DA, Beeche A, Borgemeister C, Gottlieb M, et al. Ecology: A Prerequisite for Malaria Elimination and Eradication. PLoS Medicine 2012;7(8):e1000303. doi:10.1371/journal.pmed.
Delacollette C, D'Souza C, Christophel E, Thimasarn K, Abdur R, Bell D, et al. Malaria trends and challenges in the Greater Mekong Subregion. Southeast Asian Journal of Tropical Medicine and Public Health 2009;40:674.
Organisation Internationale pour les Migration. Situation report on international migration in East and Southeast Asia. 2008.
Thang ND, Erhart A, Speybroeck N, Xa NX, Thanh NN, Van Ky P, et al. Long-lasting insecticidal hammocks for controlling forest malaria: a community-based trial in a rural area of central Vietnam. PloS One 2009;4:e7369.
Deparis X, Frere B, Lamizana M, N'Guessan R, Leroux F, Lefevre P, et al. Efficacy of permethrin-treated uniforms in combination with DEET topical repellent for protection of French military troops in Cote d'Ivoire. Journal of Medical Entomology 2004;41:914-21.
Wallace MR, Sharp TW, Smoak B, Iriye C, Rozmajzl P, Thornton SA, et al. Malaria among United States troops in Somalia. The American Journal of Medicine 1996;100:49-55.
Collins WE. Plasmodium knowlesi: A Malaria Parasite of Monkeys and Humans. Annual Review of Entomology 2012;57.
Baird JK. Malaria zoonoses. Travel Med and Infect Dis 2009;7:269-77.
Sandosham AA, Thomas V. Malariology: with special reference to Malaya: Coronet Books; 1983.
Rahman WA, Che'Rus A, Ahmad AH. Malaria and Anophelesmosquitos in Malaysia. Southeast Asian Journal of Tropical Medicine and Public Health 1997;28:599.
Vythilingam I, Foo LC, Chiang GL, Chan ST, Eng KL, Mahadevan S, et al. The impact of permethrin impregnated bednets on the malaria vector Anopheles maculatus(Diptera: Culicidae) in aboriginal villages of Pos Betau Pahang, Malaysia. Southeast Asian Journal of Tropical Medicine and Public Health 1995;26:354-8.
Chang MS, Hii J, Buttner P, Mansoor F. Changes in abundance and behaviour of vector mosquitoes induced by land use during the development of an oil palm plantation in Sarawak. Trans Roy Soc Trop Med Hyg 1997;91:382-6.
Hii JLK, Kan S, Vun YS, Chin KF, Lye MS, Mak JW, et al. Anopheles flavirostrisincriminated as a vector of malaria and Bancroftian filariasis in Banggi Island, Sabah, Malaysia. Transactions of the Royal Society of Tropical Medicine and Hygiene 1985;79:677-80.
Chareonviriyaphap T, Bangs MJ, Ratanatham S. Status of malaria in Thailand. Southeast Asian Journal of Tropical Medicine and Public Health 2000;31:225-37.
APMEN. Vietnam joins Asia Pacific Malaria Elimination Network (APMEN). http://apmenorg/storage/newsmedia/Vietnam assessed 18 Sept 2012.