Journal of Biosafety and Biosecurity 3 (2021) 125–130

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Research Article

Potential geographical distribution of Anopheles gambiae worldwide

under climate change
Chao Li a,b, Yuan Gao b, Zhe Zhao b,c, Delong Ma a,b, Ruobing Zhou b, Jun Wang b, Qinfeng Zhang a,⇑,
Qiyong Liu b,c,⇑
a
School of Public Health, Shandong First Medical University, Taian 271016, China
b
State Key Laboratory of Infectious Disease Prevention and Control, Collaborative Innovation Center for Diagnosis and Treatment of Infectious Diseases, National Institute for
Communicable Disease Control and Prevention, Chinese Center for Disease Control and Prevention (China CDC), Beijing 102206, China
c
Shandong University Climate Change and Health Center, School of Public Health, Shandong University, Jinan 250012, China

a r t i c l e i n f o a b s t r a c t

Article history: Background: Anopheles gambiae (An. gambiae) is considered the most effective malaria vector worldwide
Received 13 August 2021 and is widely distributed in Africa. The purpose of this study is to determine the potential impact of cli-
Accepted 18 August 2021 mate change on An. gambiae and predict the present and future potential suitable habitat globally.
Methods: In this study, environmental variables, global occurrence data of An. gambiae, and the maxi-
mum entropy model (MaxEnt) were used to evaluate the contribution of environmental factors and pre-
Keywords: dict the habitat suitability of An. gambiae under the current and future scenarios.
Anopheles gambiae
Results: Among all environmental variables, isothermality (Bio3, 34.5%) contributed the most to An. gam-
Potential distribution
Future climate scenarios
biae distribution. Under current climate conditions, the potential suitable areas for An. gambiae are
mainly located near the equator (approximately 30°N-30°S), with a total area of 16.58 million km2,
including central and northern South America, a fraction of areas near the equator of North America, cen-
tral and southern Africa, some tropical regions of southern Asia, and small areas of Oceania. The areas of
potential suitable habitats would be reduced to varying degrees in future climate scenarios.
Conclusions: Potential suitable habitats for An. gambiae may not be limited to Africa. Necessary surveil-
lance and preventive measures should be undertaken in high-risk regions, including those outside
Africa, to monitor and control the spread of An. gambiae.
Ó 2021 Published by Elsevier B.V. on behalf of KeAi Communications Co., Ltd. This is an open access article
under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

1. Introduction
Global warming is an indisputable phenomenon; human activ-
ities were estimated to have caused approximately 1.0 °C of global
warming above pre-industrial levels, with a likely range of 0.8 °C to
1.2 °C. Global warming is likely to increase by 1.5 °C between 2030
and 2052 if it continues to increase at the current rate1, posing a
serious threat to human health and that of the environment. The
Abbreviations: SDM, species distribution models; MaxEnt, maximum entropy;
GBIF, Global Biodiversity Information Facility; SSPs, Shared Socioeconomic Path-
ways; AIC, Akaike information criterion; RM, regularization multiplier.
⇑ Corresponding authors at: State Key Laboratory of Infectious Disease Prevention
and Control, Collaborative Innovation Center for Diagnosis and Treatment of
Infectious Diseases, National Institute for Communicable Disease Control and
Prevention, Chinese Center for Disease Control and Prevention (China CDC), Beijing
102206, China (Q. Liu).
E-mail addresses: qfzhang@sdfmu.edu.cn (Q. Zhang), liuqiyong@icdc.cn (Q. Liu).
rise in global temperature will also change the spatial distribution
of disease vectors, increasing the risk of disease2. Globally, an esti-
mate of 229 million malaria cases in 2019 in 87 malaria-endemic
countries was provided in the 2020 World Health Organization
malaria report. At the Global Technical Strategy for Malaria
2016–2030 baseline of 2015, there were 218 million estimated
malaria cases. The Global Technical Strategy for Malaria 2016–
2030 was adopted by the World Health Assembly in May 2015
and provides a comprehensive framework to guide countries in
their efforts toward malaria elimination. The strategy sets a target
of reducing global malaria incidence and mortality rates by at least
90% by 20303. Therefore, predicting the impact of climate change
on the distribution of potentially suitable areas of malaria vectors
is important.
The transmission of malaria mainly depends on the vector
Anopheles. Anopheles gambiae (An. gambiae) is an important mos-
quito vector for Plasmodium falciparum malaria, responsible for
high morbidity and mortality rates in Africa, with approximately

https://doi.org/10.1016/j.jobb.2021.08.004
2588-9338/Ó 2021 Published by Elsevier B.V. on behalf of KeAi Communications Co., Ltd.
This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
C. Li, Y. Gao, Z. Zhao et al.
384,000 deaths in 20194,5. Climate change significantly affected An.
gambiae distribution because its entire life cycle is affected, start-
ing with the hatching of eggs. The number of eggs laid was highest
when adults were kept at 27 °C6, and the optimum temperature for
egg hatching ranged from 24 °C to 30 °C, irrespective of incubation
period7. Increases in temperature increased the developmental
rates of first instar larvae and pupae but decreased developmental
rates of fourth instar larvae7.
Various species distribution models (SDMs), such as genetic
algorithms for rule set production (GARP), CLIMEX, and maximum
entropy (MaxEnt), have been used to assess ecological require-
ments, ecological responses, and suitable distribution areas8–13.
Among SDMs, MaxEnt is widely used and shows excellent perfor-
mance with small sample sizes relative to other modeling meth-
ods14,15. For example, Godwin et al. used MaxEnt to predict the
potential future distribution of major transmitters of malaria in
Nigeria under low- and high-emission scenarios; they found that
An. gambiae species were projected to experience a large shift in
the potential range and population with increased distribution
density16.
Many studies have explored the impact of climatic variables on
the ecology of An. gambiae16,17, but few studies have investigated
the potential distribution of An. gambiae in the context of climate
change worldwide. For example, An. gambiae are mainly dis-
tributed in central and southern Africa, but where will they spread
between 2040 and 2060? Therefore, using the MaxEnt model to
reasonably predict the potential habitat of An. gambiae, the main
vector of P. falciparum malaria, would contribute to the literature.
In our study, we aimed to explore the main ecological factors asso-
ciated with the distribution of An. gambiae and predict potentially
suitable habitats for An. gambiae by using various scenarios that
model the nature and trends of changes in current and future cli-
mates. The results provide a scientific basis for the effective mon-
itoring and control of An. gambiae in different world regions.
2. Methods
2.1. Occurrence data
Global occurrence data for the An. gambiae were mainly
acquired from the Global Biodiversity Information Facility (GBIF.
org [November 14, 2020] GBIF Occurrence Download https://doi.
org/10.15468/dl.wnvcmr). GBIF is an international network and
data infrastructure funded by governments worldwide; many
institutions worldwide provide its data, mainly the major distribu-
tion and scientific nomenclature of plants, animals, fungi, and
microorganisms. This helps scientists, researchers, and other orga-
nizations apply this data to hundreds of peer-reviewed publica-
tions and policy documents each year. The themes of these
analyses are often related to the effects of climate change, the
spread of invasive and exotic pests, protected areas and protected
areas, food security, and human health. We also conducted a
detailed literature search to ensure that the occurrence data of
An. gambiae were not missed, using the PubMed, Google Scholar,
and Web of Science databases. To avoid the over-fitting of the data
and being limited by the accuracy of the climate data grid, we
deleted duplicate records and records without latitude and
longitude.
Because the occurrence data were affected by the accessibility
of distribution areas and the bias of manual research, we per-
formed the following steps to reduce the problems associated with
spatial sampling biases18. First, we imported the occurrence points
into ArcMap to compare the Euclidean distances among all distri-
bution points. If two localities were within a 5 km  5 km grid,
one was randomly reserved for the subsequent analysis. This result
126
Journal of Biosafety and Biosecurity 3 (2021) 125–130
ensured that no two localities were within a 5 km  5 km grid of
one another, yielding 289 records in the final dataset. Second,
according to the requirements of ‘‘Samples” of MaxEnt (ver-
sion3.4.1) (http://www.cs.princeton.edu/wschapire/Maxent/), the
data were sorted and converted into CSV files. An observation
map was developed using ArcGIS (version 10.6) (Esri, Redlands,
California, USA) with a total of 289 records worldwide (Fig. 1).
2.2. Environmental variables and processing
All environmental data from 1971 to 2000 were obtained from
WorldClim Version 2.1 (https://www.worldclim.org/) to predict
the potential suitable habitat of An. gambiae during current and
future periods. For future climate scenarios, we used BCC-CSM2-
MR climate change modeling data under the Shared Socioeconomic
Pathways (SSPs) 126, 245, 370, and 585 scenarios released by Cou-
pled Model Intercomparison Project Phase 6. BCC-CSM2-MR is a
new, improved version of the medium-resolution climate system
model developed by the Chinese National Climate Center;
improvements are in the simulation ability of the East Asian sum-
mer monsoon, East Asian precipitation, and Madden-Julian Oscilla-
tion. These environmental parameters were pre-processed to a
general spatial resolution of 5 arcminutes with an area of 81 km2
in each grid unit.
Fifty-six initial environmental variables—19 bioclimatic vari-
ables, 36 months’ climatic variables, and 1 terrain (elevation) vari-
able—that may affect the distribution of An. gambiae were used to
establish the model. To avoid the problem of data over-fitting, we
used Pearson correlation analysis to explore the correlation among
all environmental factors by using R (version 4.0.3; https://www.r-
project.org/) software. Variables with an absolute value of the cor-
relation coefficient < 0
8 were retained directly. Variables with the
absolute value of the correlation coefficient > 0
8 were considered
highly correlated. Next, according to the results of model fitting,
the variable with the highest percent contribution was included
in the model. Of the 56 variables, 13 were selected as the forecast
variables (Table 1).
2.3. Modeling procedure
The maximum entropy model was used to predict the potential
suitable habitat for An. gambiae for the current and future periods.
In this approach, identifying the correct weighting factors or priors
is important before maximizing the entropy function subject to
constraints19. In this study, we did not use the default setting
parameters of the MaxEnt system to run the model, because opti-
mizing or adjusting the parameter settings could improve the out-
put quality of MaxEnt 20,21. The ENMeval data package in R was
used for optimal parameter selection by testing the Akaike infor-
mation criterion correction (AIC) values. AIC is a standard to mea-
sure the goodness of fit of a statistical model and generally
prioritizes the parameters with small AIC values for simulation.
Parameter values corresponding to the minimum AIC value were
selected as the parameter values used in this study. The corre-
sponding feature combination parameters were linear [L], quadra-
tic [Q], and hinge [H], and the corresponding regularization
multiplier (RM) value was 3
5. In addition, we introduced the
occurrence points for An. gambiae and selected environmental fac-
tors into the MaxEnt model, randomly selected 75% of occurrence
points to build the model, and tested the remaining 25% of occur-
rence points. In the model, jackknife tests were used to analyze the
contribution rate and importance of variables, and the area under
the curve values was used to judge the excellent accuracy of the
model22,23. The final potential distribution map was presented
using ArcGIS software. Values, which ranged from 0 to 1, were
C. Li, Y. Gao, Z. Zhao et al. Journal of Biosafety and Biosecurity 3 (2021) 125–130

Fig. 1. Original occurrence records for Anopheles gambiae.

Table 1 and Nicaragua), central South America (including Brazil), southern

Relative percentages of bioclimatic variables used in MaxEnt to model the current and Africa (including Angola, Zambia, Zimbabwe, and South Africa),
future habitat suitability of Anopheles gambiae.
central India and Laos in Asia, and the western and northern coast
Abbreviation Description % of Australia (Fig. 2).
Contribution
Bio3 Isothermality (BIO2/BIO7) (100) 34.5
Bio11 Mean Temperature of Coldest Quarter (°C) 34.3 3.2. Prediction of the spread trend of An. gambiae under the SSP
Bio16 Precipitation of Wettest Quarter (mm) 12.6 scenario
Prec1 Precipitation in January 4.4
Bio15 Precipitation Seasonality (Coefficient of 3.1
Variation) 3.2.1. Secular trend of the SSP scenario
Tmin3 Minimum temperature in March 3 By incorporating the climate data under future SSP scenarios
Tmax12 Maximum temperature in December 2.1 and the existing occurrence data into the MaxEnt model, we pre-
Prec11 Precipitation in November 2.1 dicted the distribution range of An. gambiae for the periods
Tmax8 Maximum temperature in August 1.5
Prec10 Precipitation in October 0.7
2021–2040, 2041–2060, 2061–2080, and 2081–2100. The results
Ele Elevation(m) 0.6 showed that the total suitable area presented a decreasing trend
Prec7 Precipitation in July 0.6 in different periods and scenarios in the future. For example, under
Bio19 Precipitation of Coldest Quarter (mm) 0.3 the SSP126 climate scenario from 2041 to 2060, the potential suit-
BIO2 = Mean Diurnal Range (Mean of monthly (max temp - min temp)), able area was 13
10 million km2, which was 3
48 million km2 less
BIO7 = Temperature Annual Range (BIO5 - BIO6), BIO5 = Max Temperature of than the current suitable area (Table 2 and Figs. 3 and 4).
Warmest Month, BIO6 = Min Temperature of Coldest Month.

3.2.2. Secular trend of An. gambiae distribution for different periods in

regrouped with a span of 0
2 to show potential suitable habitats.
The values suggested a ‘‘highly suitable” (>0
6), ‘‘moderately suit-
able” (0
4–0
6), ‘‘generally suitable” (0
2–0
4), and ‘‘not suitable”
(<0
2).
3. Results
3.1. Potential distribution of An. gambiae under current conditions
the same scenario
In the same scenario and different periods, the potentially suit-
able area of An. gambiae showed a downward trend. Under the cli-
mate scenarios of SSP126, SSP245, and SSP370, the total suitable
area of An. gambiae would continue to reduce from 2021 to 2080
and rebound from 2081 to 2100. Under the climate scenario of
SSP585, the total suitable area of An. gambiae showed a continuous
downward trend from 2021 to 2100 (Table 2 and Figs. 3 and 4).

Under current climate conditions, potential suitable areas for
An. gambiae are mainly located near the equator (approximately
30°N 30°S), with a total area of 16
58 million km2, including cen-
tral and northern South America, a fraction of areas near the equa-
tor of North America, central and southern Africa, some tropical
regions of southern Asia, and small areas of Oceania. High and
moderately suitable areas were mainly distributed in central and
southern Africa (including Gambia, Ghana, Nigeria, Central Africa,
Sudan, Madagascar, Mozambique); southern and southeastern Asia
(including the southwest coast of India, Thailand, Cambodia, and
Vietnam); the westernmost, easternmost, and northern parts of
South America (eastern Brazil, Ecuador, Venezuela); and the north-
ern coast of Australia. Low suitable areas were mainly distributed
along the southwest coast of North America (including Mexico
127
3.2.3. Secular trend of An. gambiae distribution in different scenarios
in the same period
Compared with the climatic conditions from 1971 to 2000, the
distribution range of An. gambiae decreased in different scenarios.
In 2061–2080 and 2081–2100, the potentially suitable areas of
An. gambiae decreased with an increase in the SSP scenario level.
This finding shows that under the global warming trend without
climate policy intervention, the suitable area of An. gambiae will
continue to decrease. Under different scenarios in the same period,
potential suitable areas for An. gambiae were reduced to different
degrees. For example, under the SSP126 scenario in 2081–2100,
the area was reduced by approximately 2
11 million km2 (13%),
and under the SSP585 scenario, the area was reduced by 7
93 mil-
lion km2 (48%) (Table 2 and Figs. 3 and 4).
C. Li, Y. Gao, Z. Zhao et al. Journal of Biosafety and Biosecurity 3 (2021) 125–130

Fig. 2. Potential distributions of Anopheles gambiae under current climate conditions.

Table 2
Current and future potential suitable area of An. gambiae all over the world under different climatic scenarios (*106km2).

Climate scenario Period Lowly suitable Moderately suitable Highly suitable Total area Area change Area change ratio (%)
current 1970-2000 8.66 3.95 3.97 16.58
ssp126 2021-2040 8.10 3.69 3.32 15.11 1.47 9
2041-2060 7.98 3.61 2.81 14.40 2.18 13
2061-2080 6.73 3.36 3.01 13.10 3.48 21
2081-2100 7.71 3.43 3.33 14.47 2.11 13
ssp245 2021-2040 8.10 3.67 2.99 14.75 1.83 11
2041-2060 7.44 3.45 2.54 13.43 3.15 19
2061-2080 6.93 2.94 2.14 12.01 4.57 28
2081-2100 7.60 3.91 2.62 14.13 2.45 15
ssp370 2021-2040 8.00 3.72 3.43 15.15 1.43 9
2041-2060 6.32 2.66 2.14 11.12 5.46 33
2061-2080 6.43 2.50 1.32 10.25 6.33 38
2081-2100 7.12 3.39 2.25 12.75 3.83 23
ssp585 2021-2040 8.01 3.77 3.54 15.31 1.27 8
2041-2060 6.91 2.90 1.88 11.68 4.90 30
2061-2080 6.12 2.15 1.10 9.37 7.21 43
2081-2100 5.73 2.01 0.91 8.65 7.93 48

4. Discussion
4.1. Significance of model predictions and the contribution of major
environmental factors
Climate change and land use have led to habitat changes in
many species24–27. In addition, the frequent exchange of interna-
tional trade has resulted in alien species entering new regions
and destroying their ecological balance, causing serious losses to
society and the economy and endangering human life and health25.
For example, Aedes aegypti and Aedes albopictus are predicted to
spread globally28. As one of the most widely concerned and urgent
diseases worldwide, malaria is one of the three infectious diseases
under the United Nations Millennium Development Goals for pre-
vention and control29.
The results of the MaxEnt model showed that the main environ-
mental factors that affected the geographical distribution of An.
gambiae included Bio3 and Bio11; this finding indicates that the
appropriate temperature was crucial for the spatial distribution of
An. gambiae. Second, Bio16 is another important factor affecting
the growth and development of An. gambiae, indicating that rainfall
is closely related to the distribution of An. gambiae. These results are
consistent with those of related studies on habitat types of Anophe-
les in Africa30. Therefore, controlling the breeding of An. gambiae in
river beds and pools during rainy seasons at suitable temperatures
may be important to prevent and control the spread of malaria31.

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C. Li, Y. Gao, Z. Zhao et al. Journal of Biosafety and Biosecurity 3 (2021) 125–130

Fig. 3. Predicted area changes of An. gambiae in different periods and scenarios. A. Predicted area changes of An. gambiae in different periods B. Predicted area changes of An.
gambiae in different scenarios.

Fig. 4. Potential distributions of Anopheles gambiae under future climate conditions.

4.2. Expected changes in the distribution range
We used the MaxEnt model to predict the range of suitable
habitats of An. gambiae under different SSPs and found that the
area of potential suitable habitat would be reduced to varying
degrees in the next 80 years. The main reason for this reduction
may be that the average temperature would be higher and the
rainfall would be lower in the future than the meteorological con-
ditions at present, which may not be suitable for the growth of An.
gambiae. In addition, the uneven spatial distribution of rainfall and
the frequent occurrence of extreme weather may also affect the
distribution patterns of An. gambiae. Notably, the largest reduction
in potential suitable areas for An. gambiae would occur between
2081 and 2100 under the SSP585 scenario, with a decrease of
approximately 48%. The suitable habitat in the SSP126 scenarios
would decrease the least of the scenarios in the same period, indi-
cating the growth of An. gambiae showed a strong tendency toward
heat.
The conclusions of this study are consistent with those of a
study on the potential distribution of dominant malaria vector spe-
cies in tropical areas16,32. An estimate of the potential distribution
of Anopheles arabiensis in Africa suggests that by 2050, its habitat is
expected to be reduced by 48% to 61% under future climate scenar-
ios33, similar to our findings on An. gambiae. Prediction results for
the potential suitable habitat of An. gambiae showed that habitats
suitable for An. gambiae were not limited to Africa, and areas near

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C. Li, Y. Gao, Z. Zhao et al.
the equator (approximately 30°N-30°S) were suitable for An. gam-
biae, such as northern South America, southern Asia, and northern
Australia. Therefore, the relevant countries in these regions should
be vigilant in monitoring and controlling the invasion of An.
gambiae.
4.3. Advantages and limitations
The main advantages of this study were that the simulation
accuracy of MaxEnt software was higher than that of other soft-
ware and that we used the ENMeval data packet to optimize the
RM value. The climatic data used in this study were the latest ver-
sion obtained from WorldClim Version2
1, the climatic parameters
and units of which were modified. The prediction results have a
certain reference significance for the use of different versions of cli-
matic data.
However, this study has limitations. First, the current niche
model considers only the environmental adaptability of An. gam-
biae; thus, further studies should be conducted to include more
environmental, ecological, and socioeconomic variables, such as
altitude variables, vegetation cover, biological factors, and human
activities, which could affect An. gambiae distribution. Second, we
only use the maximum entropy model to predict the potential suit-
able habitat of An. gambiae; thus, other SDMs, such as GARP and
CLIMEX, will be considered in our next step to provide additional
evidence.
Funding
This research was funded by the National Key R&D Program of
China (grant number 2020YFC1200101), Perceptive assessment of
health risks caused by climate change, air pollution and health
co-benefits of low carbon transition in China (grant number
15LCI1) and the Emergency Response Mechanism Operation Pro-
gram, National Institute for Communicable Disease Control and
Prevention, Chinese Center for Disease Control and Prevention
(grant number 131031102000180007).
Declaration of Competing Interest
The authors declare that they have no known competing finan-
cial interests or personal relationships that could have appeared
to influence the work reported in this paper.
Acknowledgment
We thank Professor Juan Shi and Dr. Tai Gao, working at Beijing
Forestry University, for their technical assistance.
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