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 Epidemiology and Society Health Review| ESHR 
Vol. 5, No. 1, 2023, pp. 41-50 ISSN 2656-6052 (online) | 2656-1107 (print) 
      http://journal2.uad.ac.id/index.php/eshr/index                                                   eshr@ikm.uad.ac.id 

 
 

      10.26555/eshr.v5i1.7257  

 
 

41 

 
 

  

Research Article 
 
Distribution and behavior of Anopheles maculatus and its 
potential as a Malaria vector in Indonesia 
 
Riyani Setiyaningsih1, Sapto Prihasto1, Fahmay Dwi Ayuningrum1, Arif Suryo Prasetyo1, 
Mega Tyas Prihatin1, Sekar Negari1, Siti Alfiah1, Lulus Susanti1, Evi Sulistyorini1, Jery 
Cahyandaru1, Tri wibowo Ambar Garjito2 
 
1 Institute for Disease Vector and Reservoir Research and Development (IVRCRD), 

Salatiga, Central Java, Indonesia  
2 Vector-borne and Zoonotic Diseases Research Group, Research Center for Public Health 

and Nutrition, National Research and Innovation Agency, Salatiga, Indonesia 
 
* Correspondence: riyanisetia@gmail.com. Phone: +62081575529529 
 
Received 06 January 2023; Accepted 11 March 2023; Published 12 March 2023 
 

ABSTRACT 

Background: Anopheles maculatus is one of the mosquito species that has been confirmed 
as a malaria vector in Indonesia. The potential of a mosquito as a vector is influenced by its 
behavior. Information on the distribution and behavior of An. maculatus needs to be carried 
out to determine the potential for malaria transmission transmitted by the species in an area. 
The study aimed to obtain information on the distribution, behavior, and potential of 
maculatus as a malaria vector in several provinces in Indonesia. 
Method: Mosquito collection was carried out using human-landing collection, animal-baited 
trap, cattle-bait, light-trap, and resting morning. A survey of mosquitoes was carried out in 
29 provinces in Indonesia. Mosquitoes were identified for the species and detected the 
blood-sucking behavior with an Enzyme-linked immunosorbent assay and the presence of 
Plasmodium using a Polymerase Chain Reaction. 
Results: The results showed that An. maculatus was found in the Riau Islands, Lampung, 
Bangka Belitung, West Java, Central Java, East Java, and Central Sulawesi. Anopheles 
maculatus has known to suck the blood of humans and animals with a predominance of 
animals. Anopheles maculatus is also known to suck blood outdoors predominantly. In 
general, the activity of An. maculatus sucking blood begins around 18.00 in the evening. 
Central Java Province was the province with the highest density of An. maculatus 
mosquitoes, thereby increasing the potential for transmission of malaria cases. 
Conclusion: Anopheles maculatus was spread in Riau Islands, Lampung, Bangka Belitung, 
West Java, Central Java, East Java, and Central Sulawesi. This species was known to suck 
the blood of people and humans, and its blood-sucking activity starts around 18.00 in the 
evening. 

Keywords: Anopheles maculatus, Malaria, Vector malaria 



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INTRODUCTION 

Anopheles maculatus is a malaria vector species in Indonesia found in some provinces such 
as Central Java, West Java, DIY, East Java, Bali, Lampung, North Sumatra, West Sumatra, 
Riau, Jambi, and South Sumatra. 1–4 Anopheles maculatus was a malaria vector in Thailand, 
the Philippines, Malaysia 5,6, Afghanistan, Pakistan, China, and India.7  

The potential of mosquitoes as malaria vectors can be seen from their behavior. The results 
of research in Thailand and Purworejo Central Java, Indonesia; An. maculatus was known to 
suck the blood of animals and humans.5,8 The behavior of sucking mosquito blood significantly 
affects its potential as a malaria vector. Mosquitoes that have the behavior of sucking human 
blood have the opportunity to be able to transmit Plasmodium from human to human. Another 
factor that supports mosquitoes in becoming malaria vectors is their long life and resistance 
to Plasmodium entering their body.9–11  

Studies on the distribution and behavior of An. maculatus as a malaria vector need to be 
carried out considering the importance of this species as a malaria vector in several provinces 
in Indonesia. Mosquito behavior that influences its potential as a malaria vector is resting 
behavior and blood-sucking behavior. This was also because there were possible differences 
in mosquito behavior in locations because each region has different environmental conditions. 
Based on this background, this study aims to discover the development of An: maculatus' 
distribution behavior and its potential as a malaria vector in Indonesia.  

METHOD 

The research was conducted in 29 provinces in Indonesia. We selected three districts in each 
province and six sampling points in each district. Each point represented forest, non-forest, and 
coastal ecosystems near and far from settlements. Sampling was conducted for five days at 
each point, so with six points in one district, the sampling duration was 30 days.2 A list of 
sampling locations can be found in Table 1. DKI Jakarta Province was an exception because 
no forest ecosystem was found, so sampling was conducted at locations representing the DKI 
Jakarta area. The selected locations were Central Jakarta, West Jakarta, South Jakarta, East 
Jakarta, North Jakarta, and the Thousand Islands. 

 This study was cross-sectional, and the sample was taken at a particular time. The research 
sample was An. maculatus which was caught using various research methods in all provinces. 
The study population was mosquitoes caught in all provinces. The research variables were the 
number of An. maculatus responded every hour in each province, the number of An. maculatus 
mosquitoes caught in each method in all provinces, and Plasmodium sp positive in An. 
maculatus mosquitoes in each province. 

Mosquitoes were caught using the human landing collection method, livestock bait, Animal-
baited Trap (ABT), light trap, and morning resting collection. Mosquitoes caught in each method 
identified their species using a mosquito identification key.12,13 The time used to catch 
mosquitoes in each technique was different. A list of fishing times and fishing hours per hour 
can be seen in Table 2. 

 

 



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Table 1. Sampling location of An. maculatus in several provinces in Indonesia 

Number  Provinsi City/County Name 
1 Papua Biak, Merauke , Sarmi 

2 Central Java Pati,  Pekalongan, Purworejo 

3 South Sumatra Banyuasin, Lahat, Organ Komeling Ilir (OKI) 

4 Central Sulawesi Parigi Moutong, Tojo Una Una, Toli-toli 

5 Aceh Aceh Timur, Aceh Barat, Pidie 

6 West Sumatra Pesisir Selatan, Padang Pariaman, Pasaman Barat 

7 Lampung Tanggamus, Lampung Selatan, Pasawaran 

8 Bangka Belitung Bangka, Belitung, Bangka Tengah 

9 West Java Garut, Subang, Pangandaran 

10 East Java Malang, Banyuwangi, Pasuruan 

11 Banten Pandeglang. Lebak, Serang 

12 Nusa Tenggara Barat Lombok Barat, Bima, Lombok Utara 

13 Nusa Tenggara Timur Belu. Ende, Sumba Tengah 

14 West Kalimantan Sambas, Ketapang, Kayong Utara 

15 South Kalimantan Barito Kuala, Kota Baru, Tanah Laut 

16 North Sulawesi Minahasa, Kota Manado, Kota Belitung 

17 Southeast Sulawesi Muna, Konawe, Bombana 

18 Maluku Maluku Tenggara Barat, Maluku Tenggara, Kepulauan Aru 

19 North Maluku Halmahera Tengah, Halmahera Selatan, Pulau Morotai 

20 Riau Bengkalis, Dumai, Kepulauan Meranti 

21 Jambi Bungo, Sarolangun, Tanjung Jabung Barat 

22 Special Region of Yogyakarta Kulon Progo, Gunung Kidul, Bantul 

23 Bali Jembrana, Badung, Karangasem 

24 Central Kalimantan Gunungmas, Pulang Pisau, Murung Raya 

25 South Sulawesi Pangkajene Kepulauan, Bulukumba, Luwu Timur 

26 West Papua Raja Ampat, Manokwari, Fak-fak 

27 Riau Islands Kota Batam, Bintan, Lingga 

28 DKI Jakarta Central Jakarta, West Jakarta, South Jakarta, East Jakarta, 
North Jakarta, the Thousand Islands. 

29 North Kalimantan Bulungan, Nunukan, Tarakan 

 

Table 2. The duration and catchment method  of Anopheles maculatus  

Method Catch time Length of catchment 
Indoor human bait 6 am - 6 pm 40 minute 
Outdoor human bait 6 am - 6 pm 40 minute 
Animal baited trap  6 am - 6 pm 15 minute 
Cattle bait 6 am - 6 pm 15 minute 
Morning resting 6 pm - 9 pm 3 hour 

 

Anopheles maculatus found in all capture methods was prepared by separating the head and 
thorax from the abdomen and then analyzing its Plasmodium's presence using PCR. Primers 
used in detecting Plasmodium were rPLU1 (5’ - TCA AAG ATT AAG CCA TGC AAG TGA – 



Setiyaningsih et al. (Distribution and behavior of Anopheles maculatus and its potential as a Malaria vector in Indonesia) 
 

 

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44 

3’), rPLU5 (5’ - CCT GTT GTT GCC TTA AAC TCC – 3’), rPLU3 (5’ - TTT TTA TAA TAA GGA 
TAA CTA CTA CGG AAA AGC TGT – 3’), rPLU4 (5’ - TAC CCG TCA TAG CCA TGT TAG 
GCC AAT ACC – 3’) 14,15 The blood-fed or half-gravid An maculatus mosquitoes will be tested 
their blood feed using the direct-ELISA method. 

RESULTS 

Based on the results of the study showed that An. maculatus was not found in all provinces in 
Indonesia. The results of the survey in 29 provinces in Indonesia, An. maculatus was 
distributed in Riau Islands, Lampung, Bangka Belitung, West Java, Central Java, East Java, 
and Central Sulawesi (Figure 1). In all seven provinces, 141 An. maculatus mosquitoes were 
collected. The highest density of mosquitoes An. maculatus was found in Central Java 
Province. 

 

Figure 1. Distribution An. maculatus in Indonesia 

Anopheles maculatus collection in each province showed different fluctuations and 
densities. Based on the behavior of sucking blood, An. maculatus suck blood starting at 
6:00 p.m. in all provinces. However, the An. maculatus activity of sucking blood generally 
does not occur all night. An. maculatus in Central Java Province was found sucking blood 
throughout the night with peak density at 21.00-22.00. Fluctuations in mosquito density 
based on blood-sucking time in each province showed in Figure 2. 



Setiyaningsih et al. (Distribution and behavior of Anopheles maculatus and its potential as a Malaria vector in Indonesia) 
 

 

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Figure 2. Density fluctuation An.  maculatus based on blood-sucking time in Indonesia. 

The results of mosquito catching by various methods show that, generally, An. maculatus 
dominant was found to suck blood in animals (96.67%), and only a small percentage were 
found to suck people's blood outside the home (3.33% ). The distribution and percentage of 
An. maculatus behavior in sucking blood in each province can be seen in Table 3. 

Table 3. Blood-sucking behavior of Anopheles maculatus in various provinces in 
Indonesia 

Province name 
The number of mosquitoes by the method 

ABT CB IHB OHB MR 
Riau Islands  0 0 0 1 0 

Lampung 0 0 0 1 0 

Bangka Belitung  0 0 0 1 0 

West Java 0 0 0 1 0 

Central Java 68 26 0 0 0 

East Java 1 0 0 0 0 

Central Sulawesi 21 0 0 0 0 

Total 90 26 0 4 0 

The total base of blood-sucking behavior 116 4  

Percentage of mosquitoes sucking animal blood. 
 

96.67   

Percentage of mosquitoes sucking human blood.  3.33  

 

DISCUSSION  

Our study shows that An. maculatus was found in some provinces in Indonesia, for example, 
in the Riau Islands, Lampung, Bangka Belitung, West Java, Central Java, East Java, and 
Central Sulawesi. Based on previous studies, An. maculatus was known as a malaria vector 
in the provinces of Central Java, West Java, DIY, East Java, Bali, Lampung, North Sumatra, 
West Sumatra, Riau, Jambi, and South Sumatra.2 The previous study also mentioned that An. 
maculatus be a malaria vector in Laos with An. rampae, An. sawadwongporni, An. 

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Riau Islands Lampung Bangka Belitung West Java

Central Java East Java Central Sulawesi

Information: ABT: Animal-baited trap; CB: Cattle-bait; IHB: Indoor Human-bait; OHB: Outdoor human-bait; MR: Morning resting 



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46 

pseudowillmori, An. dravidicus, An. minimus, An. aconitus, An. pampanai, An. harrisoni, An. 
dirus, An. baimaii, An. nemophilou,16 Peninsula Malaysia, along with An. cracens.17 In addition, 
An. maculatus was also known as the primary malaria vector in Morong, Bataan, Philippines, 
along with An. flavirostris.18 

The discovery of An. maculatus in several provinces in Indonesia shows the potential for 
transmission and an increase in cases of malaria transmitted by An. maculatus. East Java 
Province was a malaria-receptive area because  An. maculatus was found, previously 
confirmed as a malaria vector despite receiving a malaria elimination certificate. Meanwhile, 
Riau, Lampung, Bangka Belitung, West Java, Central Java, and Sulawesi provinces can 
transmit malaria because the provinces were still found in malaria-endemic areas.19 

The potential of An. maculatus as a malaria vector can be seen from its blood-sucking and 
resting behavior. Based on the results of studies showing that An. maculatus was known to 
be dominantly found to suck the blood of livestock, but it was also found to suck human blood. 
Although it was not dominantly found to suck human blood, contact with humans allows 
malaria transmission in the local area. This was because most of the regions found by An. 
maculatus were malaria-endemic areas.19 Studies on China's Hainan Island also show An. 
maculatus was found to suck human and animal blood. Anopheles maculatus in the area was 
known to have blood-sucking behavior several times during one gonotrophic cycle. This 
condition increases the possibility of malaria transmission.20 Results of research in 
Bangladesh An. maculatus was also known to suck human and animal blood.21 Study in Laos 
An. maculatus was the dominant species whose behavior was sucking animal blood (98%).16 
Study in Pu Teuy Village, Sai Yok Region, Kanchanaburi Province, west-central Thailand, and 
Morong, Bataan Pilipina An. maculatus was also found to be predominantly sucking the blood 
of livestock compared to human blood.05,18 Study in Vietnam An. maculatus tends to be 
zoophilic and predominantly sucks the blood of buffaloes and cows.22 

The potential of An. maculatus as a vector was also influenced by its ability to survive when 
Plasmodium enters and develops in its body. Although the results showed no discovery of 
Plasmodium in the An. maculatus examination, considering that the species has been 
confirmed as a malaria vector, the potential for malaria transmission transmitted by the species 
can occur again if in the environment there are still cases of malaria. Results of studies in 
Thailand and Myanmar An. maculatus was known to be positive for Plasmodium with a 
sporozoite index of 1.1%. Anopheles maculatus was known to be the dominant species in the 
area alongside An. sawadwongporni.23 Studies in Nepal also show An. maculatus was known 
to be sporozoite positive with An. fluviatile during the spring and found in the vicinity of cages 
in the forests of Churia Hills and during the summer in the village without spraying.24 Study in 
northern Central Vietnam. An. maculatus with An. sinensis, An. aconitus, An. harrisoni, An. 
sawadwongporni, An. peditaeniatus, and An. philippinensis found protein circumsporozoite.22 

Based on the study of An. maculatus blood-sucking activity starting at 18:00 in all provinces, 
with different fluctuations. In general, blood-sucking activities do not occur throughout the 
night. In contrast to An.  maculatus in Central Java Province, it was found that blood-sucking 
activity occurred throughout the night. This will enlarge contact with humans allowing the 
transmission of malaria. The results of a study in the Philippines An. maculatus found that the 
peak of density occurred at 22.00-00.00. The opportunity of An. maculatus as a malaria vector 
was also influenced by its density. Central Java had the highest density of mosquitoes 
compared to other provinces. High density has the potential to increase the occurrence of 



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contact with humans, thereby increasing the chances of malaria transmission. The results of 
the research in Purworejo, Central Java, An. maculatus was found to be dominant compared 
to other species. Some of the species found together with An. maculatus include An. 
balabacensis (19.76%), An. vagus (11.74%), An. kochi (5.23%), An. barbirostris (3.53%), An. 
aconitus (3.40%), An. minimus (1.28%), An. flavirostris (1.00%), An. annularis (0.04%), An. 
tesselatus (0.01%), and An. koliensis (0.01%).8 Based on studies in Thailand, An. maculatus 
has a high density in the highlands, and the highest density occurs in the rainy season.25 A 
survey in Bataan Pilipina An. maculatus revealed that peak density occurs at the end of the 
dry season.18 

Based on studies showing that An. maculatus, in general, was found sucking blood outdoors. 
This behavior causes the potential for malaria transmission transmitted by An. maculatus 
tends to occur outdoors. Studies in malaria-endemic areas, umphang valley, Tak Province, 
Western Thailand An. maculatus was also found predominantly found outdoors.26 Studies in 
sub-Saharan Africa have the potential for malaria transmission to occur outdoors; this was 
because malaria vectors were found outdoors. The discovery of mosquito blood-sucking 
activities outdoors can be caused by vector control in the house with insecticidal mosquito 
nets and indoor residual spraying (IRS).27 Studies on Guinea's Bioko Island show An. gambie 
was previously found indoors after prolonged application of vector control in the home with 
insecticidal mosquito nets caused the mosquito to adapt and change its behavior of sucking 
blood outdoors.28 Studies in Sub-Saharan Africa show the application of insecticidal mosquito 
nets and the IRS can reduce cases and deaths caused by malaria. However, the application 
of vector control in the home impacts changes in vector behavior and vector types that play a 
role in malaria transmission. This will affect the control methods that should be carried out.27 
Studies in Uganda of vector control applications with the IRS and insecticidicide mosquito 
nets, in addition to decreasing vector populations and the average number of sporozoites, also 
led to changes in the type of vectors that act as malaria vectors. The mosquito that worked as 
a malaria vector before the application of vector control in the house was An. gambiae. In 
contrast, the one that acts as a vector after applying vector control was An. arabiensis. The 
species was widely found outdoors.29 Due to changes in vector behavior, vector control was 
also carried out indoors and outdoors.30 

Another factor that affects An. maculatus can play the role of a vector was the breeding places. 
In general, An. maculatus was found in ponds with clear, muddy water containing aquatic 
plants with an environmental situation of shading. An. maculatus was also found around rivers 
with slow water flow.5,6 The potential of An. maculatus is a malaria vector because 
apolipoprotein D and cathepsin D proteins were found in the salivary glands in this species.7 

CONCLUSION 

Anopheles maculatus was distributed in the Riau islands, Lampung, Bangka Belitung, West 
Java, Central Java, East Java, and Central Sulawesi. Anopheles maculatus predominantly 
sucks the blood of livestock and partially sucks the blood of people, with most of its activity 
occurring outdoors. Anopheles maculatus generally sucks blood starting around 18:00, and 
blood-sucking action typically does not happen overnight. Central Java Province had the 
highest density of mosquitoes and blood-sucking activities at night. Anopheles maculatus was 
a potential malaria vector in Central Java compared to several other provinces. 



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Acknowledgment 

The authors would like to thank the directors and all coordinators of IVRCRD who provided 
direction during the data collection until the article writing process. We also thank the 
researchers, technicians, and administrative team who have assisted in completing the 
research and all parties involved in the research process (the coordinators, team leaders, and 
enumerators). 

Declarations 

Authors' contribution 

RS was the main contributor responsible for a technical research team, supervising the 
research process, analyzing data, compiling research articles, and improving the results of 
research article revisions. SP was the main contributor to data analysis and the technical 
research team. MTP is responsible for the technical team of research and data analysis. FDA, 
ASP, and SN as technical research teams. SA, LS, and ES as a technological research and 
management tennis team. TWAG as a technical research team and research article reviewer. 
JC was an auxiliary contributor in charge of creating a distribution map of An. maculatus 

Funding statement 

The Ministry of Health of the Republic of Indonesia funded this research. Research funding 
was provided from 2015-2018 

Conflict of interest 

There was no conflict of interest in this research.  

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