Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

A new vector emerges? Aedes vittatus (Diptera: Culicidae): ecological

description and current and future potential global geographic invasion

Estefanía Mejía-Jurado1, 2; https://orcid.org/0000-0003-3105-5998

Emmanuel Echeverry-Cárdenas1; https://orcid.org/0000-0001-9827-2626

Oscar Alexander Aguirre-Obando1, 2 *; https://orcid.org/0000-0003-2480-7004

1 Escuela de Investigación en Biomatemática, Facultad de Ciencias Básicas y Tecnologías, Universidad del Quindío.

Carrera 15 Calle 12 Norte, Armenia, Colombia; emejiaj@uqvirtual.edu.co, eecheverryc@uqvirtual.edu.co, oscaragu-

irre@uniquindio.edu.co (*Correspondence)

2 Programa de Biología, Universidad del Quindío. Carrera 15 Calle 12 Norte, Armenia, Colombia.

Received 16-II-2023. Corrected 06-IX-2023. Accepted 16-V-2024.

ABSTRACT

Introduction: The Aedes vittatus mosquito is an important vector of yellow fever in Africa, with vectorial com-

petence for dengue, chikungunya and Zika. Its presence has been reported in some places in Africa, Asia, Europe,

and –recently– America. However, information on its distribution is fragmented, with limited descriptions of the

specific characteristics of its habitats.

Objective: To compile records of its occurrence, describe the ecological characteristics of its habitat and estimate

its current and future global potential invasion.

Methods: We employed both first-record data and global occurrence records to describe its habitat. Additionally,

we used an ecological niche model, specifically the MaxEnt algorithm, with bioclimatic variable layers to estimate

potential invasion areas. Since the native range of A. vittatus is unknown, we calibrated accessible areas using two

hypotheses, Africa and Asia, based on available genetic information.

Results: Regardless of its native area, A. vittatus appears to be distributed in tropical and subtropical areas in all

continents with potential to reach even currently colder climates as global climate change. It is found mainly in

tropical and urban areas, likely through transcontinental and terrestrial passive transport.

Conclusions: The mosquito can be found on all continents, ranging from sea level to 2 500 m.a.s.l., at tempera-

tures between 15 and 30 °C, and has the potential for further expansion.

Key words: Zika; Culicidae; climate change; niche modeling; tropical mosquito; vector ecology; dengue; yellow

fever; chikungunya.

RESUMEN

En español el título sería: ¿Un nuevo vector emerge? Aedes vittatus (Diptera: Culicidae):

descripción ecológica e invasión geográfica potencial mundial actual y futura

Introducción: El mosquito Aedes vittatus es un importante vector de la fiebre amarilla en África, con competen-

cia vectorial para el dengue, la chikungunya y el Zika. Se ha informado de su presencia en algunas áreas de África,

Asia, Europa y, recientemente, América. Sin embargo, la información sobre su distribución es fragmentada, con

descripciones limitadas de las características específicas de sus hábitats.

Objetivo: Recopilar registros de su presencia, describir las características ecológicas de su hábitat y estimar su

potencial de invasión actual y futuro a nivel mundial.

https://doi.org/10.15517/rev.biol.trop..v72i1.54166

INVERTEBRATE BIOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

INTRODUCTION

Aedes (Fredwardsius) vittatus (Diptera:

Culicidae) (Bigot 1861) was characterized

based on phenotypic features of adults (N =

86), larvae (N = 41), and pupae (N = 29). It

is distinguished within the Aedes genus by

several synapomorphies. These include a dark

proboscis with a median scattering of pale

yellowish scales, bilateral small patches of nar-

row white scales on the clypeus, three pairs of

distinct, small, white spots of narrow scales

on the anterior two-thirds of the scutum, a

relatively short maxillary palpus with a white-

scaled apical portion, and a white spot at the

mid-point of tibia III (in anterior view) (Bigot,

1861; Huang, 1977; Pagac et al., 2021; Reinert,

2000). However, it is currently recognized that

A. vittatus has a widespread distribution in

the tropical regions of Africa and Asia, with

some known occurrences in the Mediterra-

nean region of Europe, including France, Italy,

Portugal, and Spain, primarily due to favor-

able climate conditions (Díez-Fernández et al.,

2018; Sudeep & Shil, 2017). Similarly, this spe-

cies has recently been identified in Cuba and

the Dominican Republic (tropical region of

America) (Alarcón-Elbal et al., 2020; Pagac et

al., 2021; Pérez-Menzies et al., 2022).

The invasion of A. vittatus, as well as other

species within the same genus such as Aedes

aegypti and Aedes albopictus, into new locations

is often linked to passive transport facilitated

by large-scale anthropogenic activities, such

as transcontinental trade (Brown et al., 2011;

Medlock et al., 2012; Scholte et al., 2008), as

well as local vehicular transport, including

both public and private land cargo (Eritja et

al., 2017; Guagliardo et al., 2014). In particular,

it has been proposed that tire transport could

be a significant mechanism for the large-scale

spread of A. vittatus (Díaz-Martínez et al.,

2021). This is due to the remarkable tolerance

of Aedes and other culicid eggs to desiccation,

allowing them to remain viable at extreme

temperatures for extended periods. This allows

them to preserve their characteristics over

extensive distances or throughout various times

of the year until conditions become optimal for

hatching and subsequent development (Diniz

et al., 2017). Furthermore, their small size poses

challenges in identification and, consequently,

in vector control.

Overall, this mosquito is hematophagous,

with females feeding on the blood of goats,

cattle, humans, and sheep (Chepkorir et al.,

2018). It exhibits ecological plasticity by breed-

ing in various habitats, including rock pools,

artificial containers, tree holes, puddles, and

fresh fruit peels. It is found in forests, savan-

nas, and arid lands, displaying both sylvatic

and peri-domestic behavior (Ali et al., 2014;

Diallo et al., 2012a; Obi et al., 2022; Suganthi

et al., 2014). The presence and distribution of

this mosquito in these microhabitats are pri-

marily influenced by climate factors. Below 10

°C, conditions are unfavorable for its biological

development; between 11 and 14 °C, only a

Métodos: Utilizamos datos de primeros registros y registros de presencia a nivel global para describir su hábitat.

Además, empleamos un modelo de nicho ecológico, específicamente el algoritmo MaxEnt, con capas de variables

bioclimáticas para estimar las áreas de potencial invasión. Dado que se desconoce el área nativa de A. vittatus,

calibramos áreas accesibles bajo dos hipótesis, África y Asia, basadas en la información genética disponible.

Resultados: Independientemente de su área nativa, A. vittatus parece distribuirse en regiones tropicales y subtro-

picales en todos los continentes, con el potencial de expandirse incluso hacia climas actualmente más fríos debido

al cambio climático global. Se encuentra principalmente en áreas tropicales y urbanas, posiblemente a través del

transporte pasivo transcontinental y terrestre.

Conclusiones: El mosquito se puede encontrar en todos los continentes, desde el nivel del mar hasta altitudes de

2 500 m.s.n.m., a temperaturas que oscilan entre 15 y 30 °C, y presenta el potencial para una mayor expansión

geográfica.

Palabras clave: Zika; Culicidae; cambio climático; modelado de nicho; mosquito tropical; ecología de vectores;

chikungunya; dengue; fiebre amarilla.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

few individuals can survive, and egg and larva

mortality increases. However, between 24 and

27 °C, with humidity levels above 75 %, 75 % to

89 % of the eggs hatch. Remarkably, it exhibits

thermophilic characteristics, as evidenced by

preimaginal development occurring between

36 and 45 °C, and it displays good tolerance to

desiccation (McClelland & Green, 1970; Rob-

erts, 2004; Service, 1970).

Additionally, A. vittatus plays a crucial

role in transmitting yellow fever in rural areas

of tropical Africa (Bueno-Marí & Jiménez-

Peydró, 2010). It also possesses vector com-

petence for dengue (Angel & Joshi, 2008),

chikungunya (Diallo et al., 1999) and Zika

(Diallo et al., 2014). Similarly, A. vittatus has

the vectorial capacity to transmit arboviruses

like Japanese encephalitis, West Nile virus,

Chandipura, and Chittoor (Sudeep et al., 2020).

The diseases transmitted by species of the Aedes

genus, including those mentioned earlier, have

raised significant concerns in global public

health due to the lack of effective vaccines

for most arboviruses, with a few exceptions

that have shown limited effectiveness, such as

the dengue vaccine (Carvalho & Long, 2021;

Girard et al., 2020).

Traditional mosquito vector control meth-

ods include the use of chemical products, pri-

marily insecticides, and biological products

(Baldacchino et al., 2015; Wilson et al., 2020).

Biological methods now encompass releasing

mosquito populations infected with Wolbachia

sp. to render natural populations refractory

to the arboviruses they transmit (Caputo et

al., 2019; O’Neill, 2018), or using transgenic

mosquitoes to suppress populations and inter-

rupt the vector transmission cycles (Bellini et

al., 2013; Carvalho et al., 2015). Nevertheless,

these efforts primarily target A. aegypti and A.

albopictus, which are global vectors responsible

for the transmission of diseases such as den-

gue, Zika, and chikungunya (Weeratunga et

al., 2017). In the Americas, these diseases are

primarily transmitted by A. aegypti (Kotsakiozi

et al., 2017) and occasionally by A. albopictus,

which has evidenced possible vertical transmis-

sion of Zika, yellow fever, and dengue by this

mosquito (Alencar et al., 2021; Martins et al.,

2012; Rúa-Uribe et al., 2012). If populations of

A. aegypti or A. albopictus were controlled, A.

vittatus could become a significant vector for

these diseases.

Currently, research on A. vittatus primar-

ily centers on its vector competence for spe-

cific arboviruses in Africa (Diallo et al., 2014;

Mulwa et al., 2018) and Asia (Angel & Joshi,

2008; Sudeep et al., 2020). There is limited

and outdated research on its biology and ecol-

ogy, primarily conducted in the same regions

(Roberts, 2001; Roberts, 2004; Service, 1970).

Furthermore, there are reports of its presence in

Europe (Bueno-Marí & Jiménez-Peydró, 2010;

Eritja et al., 2018), and it has recently become

established in the Americas. Additionally, pop-

ulation genetic analysis is being conducted to

understand the origin of the A. vittatus invasion

in the Caribbean (Alarcón-Elbal et al., 2020;

Pagac et al., 2021). Hence, it is crucial to track

potential new sites of invasion by this species,

as it could complicate the epidemiological land-

scape in public health.

Ecological niche modeling (ENM) serves

as a prediction tool for identifying areas glob-

ally with suitable environmental conditions

for A. vittatus. This tool uses species presence

records and layers of bioclimatic variables,

employing a mathematical algorithm to iden-

tify geographic areas where the species could

potentially invade. This, in turn, helps identify

potential locations for arbovirus transmission

(Carvalho et al., 2017; Peterson et al., 2011).

For instance, this method has been employed

to predict the potential distribution areas of sig-

nificant mosquitoes in public health, both glob-

ally, such A. aegypti and A. albopictus (Kamal et

al., 2018) and locally, including Culex pipiens

pallens (a primary vector of lymphatic filariasis

and Japanese encephalitis) and Culex pipiens

quinquefasciatus (a primary vector of West Nile

fever and lymphatic filariasis) (Chandel et al.,

2013; Cui et al., 2013; Liu et al., 2020).

Disease-vector mosquitoes are influenced

by climatic variables such as temperature and

humidity, as they exhibit positive responses

to environmental conditions. For instance,

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

A. aegypti and A. albopictus thrive in tem-

peratures between 20 and 32 °C. Conversely,

they exhibit negative responses to temperatures

exceeding 39 °C or falling below 16 °C, which

extends their life cycle (Ezeakacha & Yee, 2019;

Marinho et al., 2016). These factors significant-

ly affect their survival, bite rates, and distribu-

tion, consequently impacting the transmission

of diseases, either positively or negatively (da

Cruz-Ferreira et al., 2017).

One approach to predict the impact of

climate change on mosquito distribution is

to use prediction models, such as the Shared

Socioeconomic Pathways (SSP), which estimate

future concentrations of greenhouse gases,

including CO2. These include SSP245, SSP370,

and SSP585, which can help identify poten-

tial future invasion areas by considering the

impact of climate change under these scenarios

(O’Neill et al., 2017; Riahi et al., 2017).

A. vittatus, along with other vectors, has

been the subject of niche modeling studies

for chikungunya virus vectors in southeast

Senegal’s Kedougou region (Richman et al.,

2018), potential distribution of Aedes genus

species in Nigeria (Omar et al., 2021), and

arbovirus vectors in Morocco (Abdelkrim et al.,

2021). Recently, a global study, which included

three mosquito species, including A. vittatus,

estimated their potential global distribution

under current and future climate conditions

(Abdelkrim et al., 2022). However, the study

did not account for the vector’s ecological

characteristics in currently registered zones.

Additionally, unlike other mosquito species

(Echeverry-Cárdenas et al., 2021; Kamal et al.,

2018), the present study did not use the native

area for calibration model. Since the native

origin of A. vittatus remains unknown, calibra-

tion of the accessible area relied on available

genetic information. Furthermore, estimations

were conducted using new models and climate

change scenarios (CMIP6) (Eyring et al., 2016).

Hence, understanding the characteristics

of its habitats and identifying potential invasion

areas, both current and future in the context of

climate change, is essential. This knowledge will

help assess its potential global epidemiological

impact and enable the inclusion of this vector

in surveillance and control strategies for local

transmission scenarios. Therefore, our objec-

tive was to describe the ecology of the new

vector, A. vittatus (Diptera: Culicidae), based

on occurrence records and estimate its global

invasion potential under two hypothetical ori-

gins, Asia, and Africa, for both the present and

the future.

MATERIALS AND METHODS

Study area: This global study identified

A. vittatus in tropical Africa, tropical Asia,

the western Mediterranean region of Europe,

and recently, in some Caribbean islands (Alar-

cón-Elbal et al., 2020; Bueno-Marí & Jimé-

nez-Peydró, 2010; Díaz-Martínez et al., 2021;

Pérez-Menzies et al., 2022; Sudeep & Shil, 2017).

Bibliographic review: We conducted a

systematic search for scientific articles report-

ing the first records of A. vittatus at the country

level. We used the following databases to iden-

tify its geographical presence: Scopus (https://

www.scopus.com/), PubMed (https://pubmed.

ncbi.nlm.nih.gov/), Springer link (https://link.

springer.com/), Science direct (https://www.

sciencedirect.com/), Google Scholar (https://

scholar.google.es/) and Google (www.google.

com). Google was employed to gather non-

scientific information, such as news, notes,

and reports. In each of the databases, keywords

were used, like “Aedes vittatus”, followed by the

Boolean operator “AND” and “first registry”

and the Boolean operator “OR” and “registries”.

To broaden the dataset of vector occurrences,

we used the keyword “Aedes vittatus” for the

general search of vector registries. We reviewed

titles and abstracts from the search results and

selected only those that met the criteria, such as

the first registry in a country or the presence of

the vector in a specific site. The search in each

database covered all dates in English, Spanish,

and Portuguese.

Registries of presence of Aedes vittatus:

We created a dataset from the results of the

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

previous section and data obtained from the

Global Biodiversity Information Facility (GBIF,

2021; https://www.gbif.org/). This dataset was

used to create a table containing all the first

registries and records reported in scientific

articles for A. vittatus in each country. The table

includes relevant information such as the con-

tinent, year of the registry, and reference. In

addition, georeferencing of this dataset served

as the foundation for estimating the global

geographic distribution and potential invasion

of A. vittatus (further details in the ‘Estima-

tion of the Geographic Distribution’ section).

Data lacking georeferencing information were

excluded from this study.

Additionally, this dataset was cleaned to

remove duplicate entries and address geospa-

tial issues (Kamal et al., 2018; Kraemer et al.,

2015). To achieve this, we used the R Studio

environment (R CoreTeam 2017) along with

the sp 1.4-1 (Pebesma et al., 2020), raster

3.4-5 (Hijmans et al., 2020a), and dismo 1.3-3

(Hijmans et al., 2020b) packages. Due to the

absence of data regarding the native area of A.

vittatus, we relied on existing genetic evidence

from the COI gene. This evidence, as presented

in the haplotype network suggested by Pagac et

al. (2021), primarily links most of the analyzed

populations to Africa (6 haplotypes) and Asia

(21 haplotypes). Haplotype networks help pro-

pose the geographic origin or ancestral lineage

of species using molecular markers (Leigh &

Bryant, 2015). Two scenarios were evaluated,

one assuming the native area of A. vittatus is in

Africa, and the other in Asia. Under the African

origin hypothesis, the occurrence dataset was

divided into two subgroups: A, which included

occurrences from Africa used for calibration,

and B, containing occurrences from the rest of

the world for validation. Similarly, under the

Asian origin hypothesis, subgroups C and D

were formed, with C including Asian occur-

rences used for calibration, and D contain-

ing occurrences from the rest of the world

for validation.

Ecological niche modeling and estima-

tion of the accessible area: To predict the

potential invasion of A. vittatus under cur-

rent and future conditions, we employed the

maximum entropy algorithm implemented in

MaxEnt v3.4.3 software (Phillips et al., 2021).

This allows us to assess the influence of envi-

ronmental variables in the estimations and

calibrate models using datasets of various sizes.

Additionally, it is suitable for predictions at

both temporal and spatial scales (Peterson et

al., 2011; Phillips et al., 2006). To calibrate the

ENM, we defined the accessible area (M) of

the species, including its occurrence records in

both continents, Africa, and Asia, following the

approach outlined by Barve et al. (2011). This

involved implementing a convex minimal poly-

gon that covered a significant portion of the

species’ distribution area, as per Rochlin et al.

(2013). Similarly, 10 000 random background

points were placed within each accessible area

for model calibration (Peterson et al., 2011).

For each continent, we employed two model-

ing methods: spatial modeling, projecting ideal

current environmental conditions for A. vit-

tatus invasion worldwide, and spatial-temporal

modeling, suggesting potential vector invasion

on a global scale considering the effects of cli-

mate change.

Climate data and ecological character-

ization: To estimate the current geographic

distribution, we used bioclimatic data from

WorldClim v2.1 (https://www.worldclim.org/),

encompassing 19 bioclimatic variables collected

from global weather stations (Fick & Hijmans,

2017). The data had a spatial resolution of 2.5

min, approximately 5 km² (Kamal et al., 2018;

Kraemer et al., 2015; Medley, 2010). These

variables underwent a Variance Inflation Factor

(VIF) analysis in R Studio using the raster 3.4-5

(Hijmans et al., 2020a), rgdal 1.5-23 (Bivand

et al., 2021), and usdm 1.1-18 (Naimi, 2017)

packages to reduce multicollinearity (Pradhan,

2016). The resulting variable layers were then

used to estimate the species potential invasion

under current and future conditions.

To estimate the future invasion of A. vitta-

tus in the context of climate change, we used the

selected variables from WorldClim v2.1 under

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

the SSP245, SSP370, and SSP585 scenarios for

the periods 2041-2060 and 2081-2100. This

was performed using Global Circulation Mod-

els (GCMs), as they offer a flexible and robust

framework to evaluate the statistical signifi-

cance and importance of variables, providing

a straightforward and solid interpretation of

the model (Flato et al., 2013; Guisan et al.,

2002; Marengo et al., 2014). We used the fol-

lowing models: GISS‒E2‒1‒H, MIROC‒ES2L,

MPI‒ESM1‒2‒HR, MRI‒ESM2‒0 (Hausfather

et al., 2022), IPSL‒CM6A‒LR, and UKESM1‒0‒

LL (Flynn & Mauritsen, 2020). These models

were selected based on their values of ECS

(2.5-4 °C) and TCR (1.4-2.2 °C) to repre-

sent different climate sensitivities. All selected

variable layers were adjusted to match the

defined accessible area (M) and were used to

construct invasion scenarios for the specified

climate change periods.

To characterize the habitats where A. vit-

tatus is currently found, we extracted tem-

perature (mean, standard deviation, minimum,

and maximum) and precipitation (mean, stan-

dard deviation, minimum, and maximum) data

from the WorldClim v2.1 dataset (https://www.

worldclim.org/). These variables are crucial for

understanding the mosquito vectors’ life cycle

(Echeverry-Cárdenas et al., 2021; Leyton et al.,

2020; Valencia-Marín et al., 2020). We obtained

and extracted elevation data from WorldClim

v2.1 (https://www.worldclim.org/) and land

coverage data from Diva-gis (https://www.diva-

gis.org/). We determined the type of urbaniza-

tion through visual exploration of occurrences

using QGIS v3.16 with the available Google

Hybrid layer. Urban occurrences were classi-

fied as those located within urban areas, while

those in urbanization perimeters or immedi-

ate vicinity were considered peri-urban areas.

Occurrences without human establishments

nearby were categorized as rural areas. Addi-

tionally, we obtained sea route data (Benden,

2022) and overlaid it on the current invasion

potential to assess the potential influence of

transcontinental freight transportation routes

on the species’ spread.

Estimation of the geographic distribu-

tion: To estimate the potential geographic dis-

tribution of A. vittatus, we considered four

contexts that accounted for latitudinal and

longitudinal variations. For model calibration,

we used subgroups A and C with environmen-

tal variable layers under current conditions

confined to accessible areas. Subsequently, we

estimated the potential effects of climate change

in areas of potential A. vittatus invasion for the

periods 2041-2060 and 2081-2100 under the

SSP245, SSP370, and SSP585 emission path-

ways for each respective period.

We conducted all estimations using the

maximum entropy algorithm in MaxEnt soft-

ware v.3.4.3 (Phillips et al., 2021). Each esti-

mation consisted of ten replicates with 1 000

iterations, using the logistic output format.

Parameters ‘Do Clamping’ and ‘Extrapola-

tion’ were deactivated for current and future

estimations to prevent ecological variable

extrapolations (non-analogous climates) (Ech-

everry-Cárdenas et al., 2021; Kamal et al.,

2018). To represent each emission scenario

under future conditions, we calculated the

mean between the predictions generated by

each selected GCM, resulting in a final map

(Abdelkrim et al., 2022). We reclassified all

resulting maps into a binary format, distin-

guishing potential invasion areas based on a

threshold corresponding to the lowest environ-

mental suitability value associated with known

presence records. We considered an omission

value (E) of 0.05 (Jiménez-García & Peterson,

2019; Liu et al., 2005; Marques et al., 2020).

The potential extension area was quantified

in all scenarios.

Validation of the model: We validated

the models only under current conditions, as

the behavior of A. vittatus under future climate

change scenarios is unknown. We used the

ntbox packages (Osorio-Olvera et al., 2020) in

R Studio to estimate the area under the curve

(AUC), with metric values between 0.7 to 0.9

indicating good models and values > 0.9 con-

sidered excellent (Peterson et al., 2011). To fur-

ther assess model performance, we determined

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

the significance level of the AUC using partial

analysis of Receiver Operating Characteristics

(ROC-partial). This analysis utilized the B and

D subsets of A. vittatus presence records (Lobo

et al., 2008). This analysis aimed to address AUC

deficiencies (Peterson et al., 2008) by adjusting

the parameter to 0.05 per 1 000 iterations. To

evaluate the model’s significance, we consid-

ered AUC values with P < 0.05, indicating that

the predictions were statistically superior to

those from a random model (Echeverry-Cárde-

nas et al., 2021; Peterson et al., 2008).

RESULTS

The systematic search resulted in over 1 300

results, with 63 meeting the search criteria.

Among these, seven articles reported the first

registries of A. vittatus, while the remaining

110 registries were from various locations in

Africa, Asia, Europe, and America (Table 1; Fig.

1A). The search on the GBIF platform initially

retrieved 395 occurrences, which were reduced

to 94 after purging. Therefore, the final dataset

from the systematic search and GBIF platform

comprises 212 occurrences (Fig. 1A).

Table 2 lists the selected variables for cali-

brating the ENM, all of which are important in

the biological cycle of A. vittatus. The supple-

mentary material (SMT1) contains the ecologi-

cal characterization data, which indicates that

A. vittatus is mainly found in urban zones (42.4

%), followed by rural (39.6 %) and peri-urban

(17.9 %) zones (Fig. 1B). In these zones, the

Table 1

Records of A. vittatus worldwide were obtained from a bibliographic review. The data are presented chronologically, from

oldest to most recent.

Continent Country Year Reference

Europe France 1861 (Bigot, 1861)*

Africa Morocco 1916 (Gaud, 1947)*

Europe Spain 1925 (Gil-Collado, 1930)*

Europe Spain 1927 (Gil-Collado, 1930)

Europe Spain 1929 (Gil-Collado, 1930)

Asia Bangladesh 1949 (Nasir-ud-din, 1952)

Africa Tunisia 1952 (Vermeil, 1953)*

Europe France 1952 (Hamon & Remmert, 1952)

Europe Italy 1960 (Coluzzi, 1961)*

Europe France 1960 (Callot, 1962)

Asia Pakistan 1960 (Qutubuddin, 1960)

Africa Zimbabwe 1967 (Freyvogel & McClelland, 1969)

Africa Senegal 1972 (Diallo et al., 1999)

Africa Uganda N.A (Mukwaya, 1974)

Africa Nigeria 1973 (Service, 1974)

Africa Nigeria N.A (Stafford, 1981)

Africa Nigeria 1981 (Irving-Bell et al., 1987)

Asia Pakistan 1993 (Suleman & Khan, 1993)

Asia India 1997 (Patel, 2016)

Asia India 1998 (Rajavel et al., 2005)

Asia Oman 2000 (Roberts, 2004)

Asia Saudi Arabia 2000 (Miller et al., 2002)

Asia Pakistan 2000 (Ilahi & Suleman, 2013)

Asia India 2006 (Angel & Joshi, 2008)

Africa Central African Republic 2006 (Ngoagouni et al., 2012)

Europe Spain 2007 (Bueno-Marí, 2010)

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

Asia India 2008 (Kumari et al., 2011)

Africa Senegal 2009 (Diagne et al., 2014)

Asia Saudi Arabia 2009 (Al Ashry et al., 2014)

Africa Nigeria 2010 (Idowu et al., 2012)

Asia Saudi Arabia 2010 (Alikhan et al., 2014)

Africa Senegal 2011 (Diallo et al., 2014)

Africa Nigeria 2011 (Adeleke et al., 2013)

Asia Iran 2011 (Nasirian et al., 2014)

Africa Ghana 2012 (Suzuki et al., 2016)

Africa Ghana 2012 (Esena et al., 2013)

Asia Pakistan 2013 (Ali et al., 2013)

Africa Ivory Coast 2013 (Zahouli et al., 2017b)

Africa Ivory Coast 2014 (Zahouli et al., 2017a)

Asia Pakistan 2014 (Khan et al., 2015)

Africa South Africa 2014 (Guarido et al., 2021)

Africa South Africa 2014 (Johnson et al., 2020)

Europe Spain 2015 (Díez-Fernández et al., 2018)

Africa Kenya 2015 (Chepkorir et al., 2018)

Africa Ghana 2015 (Captain-Esoah et al., 2020)

Europe Spain 2017 (Eritja et al., 2018)

Africa Kenya 2017 (Mulwa et al., 2018)

Africa Nigeria 2017 (Afolabi et al., 2019)

Asia United Arab Emirates 2017 (Roberts, 2020)

Asia Sri Lanka 2017 (Chandrasiri et al., 2019)

Africa Ethiopia 2017 (Ferede et al., 2018)

Africa South Africa 2017 (Omar et al., 2021)

Africa Benin 2017 (Anges et al., 2018)

Africa Sudan 2017 (Eshag et al., 2019)

Africa Tunisia 2017 (Ben Ayed et al., 2019)

Asia India 2018 (Singh et al., 2019)

Asia India 2018 (Sudeep et al., 2020)

Africa Ghana 2018 (Joannides et al., 2021)

Asia Sri Lanka 2018 (Surendran et al., 2021)

America Dominican Republic 2019 (Alarcón-Elbal et al., 2020)*

America Cuba 2019 (Pagac et al., 2021)*

America Cuba 2020 (Díaz-Martínez et al., 2021)

Asia India 2020 (Reegan et al., 2020)

* First registries; N.A Year not available.

mosquito is found in various habitats, includ-

ing tropical forests (37.2 %), tropical grass-

lands (30.1 %), deserts (16.0 %), Mediterranean

forests (9.4 %), lakes (2.8 %), high-altitude

grasslands (1.8 %), temperate forests (0.9 %),

mangroves (0.9 %), and temperate grasslands

(0.5 %). Based on global registries collected and

with a 30-year data history for temperature and

precipitation variables, on average A. vittatus in

Africa is found at temperatures of 25.4 °C ± 2.4

(min 9.7-33.3 max), precipitations up to 100.9

mm ± 85.1 (min 0.0-466.0 max), and altitudes

of 451.9 m ± 359.9 (min 4-1 656 max). In Asia,

the mosquito is found at temperatures averag-

ing 25.9 °C ± 3.0 (min 6.5-35.9 max), with

precipitation levels averaging 67.6 mm ± 57.4

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

(min 0-1 594 max), and altitudes around 415.6

m ± 521.4 (min 2-2 403 max). In Europe, the

conditions are characterized by temperatures

averaging 15.4 °C ± 5.6 (min 1.4-27 max),

precipitation averaging 74.6 mm ± 58.1 (min

2-324 max), and altitudes around 357.9 m ±

376.1 (min 19-1 497 max). In the Americas,

A. vittatus typically experiences temperatures

averaging 24.7 °C ± 1.6 (min 20.2-27.5 max),

precipitation averaging 42.3 mm ± 15.4 (min

25-222 max), and altitudes around 263.3 m ±

317.9 (min 64-630 max). Furthermore, a Krus-

kal-Wallis analysis determined that between

altitude and zone type no statistically signifi-

cant differences exist (chi-squared= 2.2375,

df= 2, P = 0.3267). In turn, in any of the three

zones, A. vittatus has been recorded principally

between 100 and 500 m approximately, never-

theless, it has been registered in altitudes up to

1 600 m in urban zones and > 2 000 m in peri-

urban and rural zones (Fig. 2).

Fig. 1. World map showing records of A. vittatus occurrence. A. Zones where it is found based on the occurrences. B. In both

maps the isoclines are presented. The isoclines show the tropical and subtropical regions.

Table 2

Climatic variables selected for the ENM of A. vittatus in

the world.

Variable Unit

Rainfall seasonality %

Isothermality %

Annual precipitation mm

Precipitation of the wettest month mm

Precipitation of the driest quarter mm

Mean annual temperature °C

Mean diurnal range °C

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

The estimations under the current con-

text, for both hypotheses of native areas, Afri-

ca (AUC= 0.90 and ROC partial P = 0.00)

and Asia (AUC= 0.83 and ROC partial P =

0.00), suggest that both models are suitable

for making predictions. The invasion scenar-

ios obtained in both estimations indicate that

A. vittatus would primarily inhabit tropical

regions rather than subtropical ones. These

models predict its potential presence in Ocea-

nia, North America, and South America, where

its presence has not been reported to date. In

these estimations, A. vittatus could be found at

altitudes of up to 2 900 m in Africa and Europe

and 3 900 m in Asia. Conversely, for the African

hypothesis, it is estimated that it could inhabit

areas up to 3 900 m in America and 900 m

in Oceania, while for the Asian hypothesis, it

could be found up to 4 900 m in America and

749 m in Oceania.

In the African hypothesis, which assumes

Africa as the native area, the proposed inva-

sion encompasses a smaller area (in square

kilometers) and fewer sites compared to the

Asian hypothesis, where Asia is considered the

native area (Table 3). In the African continent,

A. vittatus could be found in 96 % of its total

extent, spanning across 52 countries, with a

Fig. 2. Box diagram showing the results of the Kruskal-Wallis test for the global registries of A. vittatus with respect to altitude

and type of zone.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

more scattered distribution in the northern and

southern regions of the continent. In contrast,

the Asian hypothesis suggests a distribution that

covers regions in every country on the African

continent. In the Asian continent, the African

hypothesis covers 39 % of the continent (31

countries), while the Asian hypothesis covers

52 % (27 countries) with a greater distribution

towards the western regions. It is noteworthy

that both hypotheses for the Asian invasion

exhibit similarities and include regions such as

the Philippine islands, Indonesia, Japan, Malay-

sia, and Taiwan. In the European continent, the

African hypothesis encompasses suitable areas

in 91 % of the continent (45 countries). In con-

trast, the Asian hypothesis concentrates its dis-

tribution primarily in the Mediterranean region

and eastern Europe, covering approximately 86

% of the continent. Additionally, suitable areas

were identified in Iceland, which is not the case

in the African hypothesis. In Oceania, accord-

ing to both hypotheses, the mosquito could

be found in much of Australia, with a lesser

presence in New Caledonia, New Zealand, and

Papua New Guinea. Furthermore, in the Amer-

ican continent, according to both hypotheses,

the distribution extends to all countries except

Surinam under the Asian hypothesis. In this

same hypothesis, the presence of the mosquito

could also be noted along the coastline of Alas-

ka (the United States) (Fig. 3).

Overall, in the future context for both

hypotheses, the invasion of A. vittatus reaches

very similar zones. The invasion consistently

tends to increase in tropical zones regardless

of the scenario and period, and this trend

becomes even more pronounced in subtropical

zones with high emission scenarios (SSP370

and SSP585). For the Asian hypothesis, it is

estimated that A. vittatus can potentially invade

colder zones, which is not the case in the

African hypothesis. In both hypotheses, for

the 2041-2060 period and across all three sce-

narios, the mosquito could be found at altitudes

of up to 3 900 m in Asia, Africa, and America.

Nevertheless, in the Asian hypothesis, it could

reach altitudes exceeding 4 000 m in America

within all three emission scenarios. In Europe,

for both hypotheses, the mosquito could reach

altitudes up to 2 900 m. In addition, in Oceania,

the maximum altitude is between 500 and 749

m. For the 2081-2100 period, under the African

hypothesis, A. vittatus could invade altitudes

> 4 000 m in America in scenarios SSP370

and SSP585. Likewise, A. vittatus can reach

altitudes > 5 000 m in Asia (China) under the

three scenarios in the Asian hypothesis, while

for the African hypothesis, this occurs only in

Table 3

Expansion of current and future potential global invasion areas for A. vittatus.

Hypothesis Period Scenario Native area (km2)Invasion area (km2)Absence area (km2)

Africa Current ‒18 693 336.5 13 082 744.5 108 343 919.0

2041-2060 SSP245 17 679 724.5 14 006 475.7 108 433 799.8

SSP370 18 019 071.4 14 279 094.5 107 821 834.2

SSP585 18 538 825.1 14 667 354.9 106 913 820.0

2081-2100 SSP245 18 538 230.5 15 239 057.0 106 342 712.5

SSP370 25 046 653.9 26 009 563.2 89 063 782.9

SSP585 23 682 315.9 26 163 646.0 90 274 038.2

Asia Current ‒ 7 123 754.9 23 307 443.3 109 688 801.8

2041-2060 SSP245 13 010 899.4 28 791 713.6 98 317 387.0

SSP370 12 609 887.7 28 476 616.3 99 033 496.0

SSP585 14 260 564.9 29 400 383.6 96 459 051.5

2081-2100 SSP245 18 498 701.5 29 065 207.7 92 556 090.9

SSP370 17 591 115.3 29 199 422.3 93 329 462.4

SSP585 17 555 140.0 28 984 905.5 93 579 954.5

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

high scenarios (SSP585). Furthermore, within

the invasion zone encompassing Greenland

and Antarctica, the mosquito could be found at

altitudes reaching up to 1 900 m.

In future estimations, under a context

of climate change, in the African hypothesis

for the first period (2041-2060), the potential

invasion of A. vittatus in the three scenarios

(SSP245, 370 and 585) was quite similar to the

current conditions. In SSP245 scenario, the

invasion area expanded compared to the inva-

sion under current conditions (Table 3). This

is evident in countries across the Americas,

including Brazil, Guyana, Suriname, French

Guiana, Honduras, El Salvador, Guatemala,

Belize, Puerto Rico, the Dominican Republic,

the United States, and Canada. However, the

potential invasion area decreased in Argen-

tina, Colombia, Ecuador, Panama, and Mexico.

Meanwhile, in Asia, it expanded in China and

Fig. 3. Model of global potential invasion by A. vittatus under current conditions, assuming A. Africa, and B. Asia as its native

regions. In both maps the principal seaports and their principal maritime trade routes are represented.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

South Korea but contracted in Saudi Arabia,

Oman, Yemen, India, Turkey, Iran, Kazakh-

stan, southwestern Russia, Burma, Iraq, and

North Korea. In the European continent, the

potential invasion area expanded in Norway,

Sweden, Poland, Ukraine, Romania, Hungary,

Slovakia, Lithuania, and the Czech Republic

but decreased in Spain. Furthermore, in Africa,

it increased in certain areas of Tanzania and

Kenya but diminished in Morocco, Algeria,

Mauritania, Mali, Niger, Chad, Sudan, Ethio-

pia, Somalia, Eritrea, the Democratic Republic

of Congo, and Madagascar. In Oceania, the

potential invasion area decreased in Australia

and New Zealand (Fig. 4A).

In SSP370 scenario, both the invasion area

and the native distribution area expanded (Table

3). The trends are observed in various countries

across different continents. In the Americas,

countries such as Colombia, Brazil, Venezu-

ela, Guyana, Suriname, Peru, Bolivia, Argen-

tina, Panama, Nicaragua, Guatemala, Belize,

the Dominican Republic, Puerto Rico, Mexico,

the United States, and Canada have seen these

changes. In Asia, we have witnessed increases

in Laos, Vietnam, Kazakhstan, Turkmenistan,

Turkey, Saudi Arabia, Pakistan, Tajikistan, and

Yemen, with decreases noted in Oman and

China. In Europe, expansion has been observed

in Sweden, Poland, Belarus, Ukraine, Romania,

and Spain. In Africa, the invasion has intensi-

fied in Algeria, Morocco, Libya, Egypt, Nigeria,

Tanzania, Kenya, Uganda, Botswana, Congo,

Zimbabwe, and South Africa, while variation

Fig. 4. A.-C. Global potential invasion model for Africa and, D.-F. Asia, during the 2041-2060 period under three climate

change scenarios (SSP245, SSP370, and SSP585).

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

is evident in Senegal, Mali, Niger, Chad, and

Sudan. Finally, Oceania has seen an increase

in Australia (Fig. 4B). In scenario SSP585, the

invasion increases (Table 3) for the sites men-

tioned, including India (Asia), diminishing in

Senegal, Mali, Niger, Chad, and Sudan (Africa)

(Fig. 4C).

For the second period (2081-2100) in sce-

nario SSP245, the potential invasion remained

similar to that of the first period, with a broader

extension observed in the Americas, including

Brazil, Guyana, Suriname, French Guiana, Peru,

Colombia, Venezuela, Argentina, Bolivia, Pan-

ama, Nicaragua, Honduras, Guatemala, Belize,

El Salvador, Mexico, Puerto Rico, the Domini-

can Republic, the United States, and Canada.

In Asia, the invasion expands in China, India,

Pakistan, Tajikistan, Kazakhstan, Afghanistan,

Turkmenistan, Turkey, Iraq, Iran, Saudi Arabia,

southwestern Russia, and South Korea, while

decreasing in Oman and Yemen. In Africa, the

potential distribution varies across the conti-

nent, particularly in Algeria, Morocco, Libya,

Egypt, Eritrea, Senegal, Mali, Niger, Chad,

Sudan, Angola, and the Democratic Republic

of Congo. In Oceania, the invasion increases in

Australia (Fig. 5A).

In scenarios SSP370 and SSP585, the inva-

sion of A. vittatus in the continents is broader

(Table 3), given that for the first scenario,

it reaches almost the entire American conti-

nent, except for Canada and some sites in the

United States and around Colombia, Venezu-

ela, Brazil, Ecuador, and Peru. Similarly, the

Fig. 5. A.-C. Global potential invasion model for Africa and, D.-F. Asia during the 2081-2100 period under three climate

change scenarios (SSP245, SSP370, and SSP585).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

invasion extends to nearly the entire African

continent, with estimates suggesting an inva-

sion into parts of the Sahara. In Europe, the

distribution remains unchanged compared to

scenario SSP245. However, in Asia, the invasion

extends into the central part of the continent

and intensifies around Iran, Saudi Arabia, and

in islands such as Malaysia and Indonesia. In

Oceania, the invasion increases in some zones

of Australia, New Zealand, and Papua New

Guinea (Fig. 5B). In scenario SSP585, the inva-

sion remains unchanged compared to scenario

SSP370; nevertheless, a decrease is projected in

certain areas of South America, southern Asia,

and Australia (Fig. 5C).

In the Asian hypothesis for the initial

period (2041-2060), the potential invasion of

A. vittatus in scenario SSP245 increases in

the southern part of Asia, particularly around

Saudi Arabia and Iran, as well as in North

and South America. However, there are varia-

tions in certain areas of Africa, such as the

Democratic Republic of Congo and countries

surrounding the Sahara (Morocco, Mauritania,

Senegal, Mali, Niger, Chad, and Sudan). In

turn, in Europe, the invasion increases in Spain,

Sweden, the United Kingdom, Italy, France,

Rumania, Ukraine, Germany, and Austria (Fig.

4D). In scenarios SSP370 and SSP585, the

distribution remains quite similar to scenario

SSP245 except for some zones where the inva-

sion diminishes for scenario SSP370, as in

America (Argentina, Mexico, the United States,

and Canada), Asia (Saudi Arabia, China, and

Afghanistan), Europe (Germany, Poland, Belar-

us, Ukraine, Sweden, and Finland), and Africa

(Mauritania, Mali and Sudan) (Fig. 4E). In sce-

nario SSP585, the invasion increases for all con-

tinents, varying in Africa in Mauritania, Mali,

Niger, Chad, and Sudan (Table 3) (Fig. 4F).

For the second period (2081-2100) across

all three emission scenarios, an invasion is pro-

jected in southern Greenland and an area of the

Antarctic, extending through South America

(Fig. 5D, Fig. 5E, Fig. 5F). In scenarios SSP245

and SSP370, the invasion expands in North

America, encompassing larger areas in Canada

and the United States, and in South America,

affecting certain regions of Brazil, Colombia,

Venezuela, Peru, Ecuador, and Argentina. Simi-

larly, an increase is projected in the Asian

continent in China, Mongolia, and southeast-

ern Russia. However, for scenario SSP370, a

decrease is expected in certain areas of Africa,

particularly around the Sahara (Burkina Faso,

Mauritania, Niger, Nigeria, Somalia, Eritrea,

Senegal, Mali, Niger, Chad, Sudan, South

Sudan) (Fig. 5D, Fig. 5E). For scenario SSP585,

a decrease is estimated in the invasion in

South America, Africa, and Australia, with an

increase in some zones of the United States, like

Alaska (Fig. 5F).

DISCUSSION

The A. vittatus mosquito is found in tropi-

cal and subtropical regions on all continents,

except Oceania. It is distributed across urban

areas (42.4 %), rural areas (39.6 %), and peri-

urban zones (17.9 %), with altitudes ranging

from 2 to 2 403 m. No global studies exist for

A. vittatus focusing on its latitudinal and altitu-

dinal distribution. Existing studies are based on

faunal surveys that indirectly address A. vittatus

in rural, peri-urban, and urban zones in Africa

(Captain-Esoah et al., 2020; Diallo et al., 2012a;

Diallo et al., 2012b) and Asia (Alikhan et al.,

2014; Angel & Joshi, 2008; Nasirian et al., 2014).

Furthermore, our current estimations indicate

the invasion of large areas in countries across

every continent. Consequently, A. vittatus can

currently be found in rural, peri-urban, and

urban zones. Various studies on other invasive

mosquito species and disease vectors suggest

that the spread and colonization of new areas

are associated with factors such as urbaniza-

tion, climate, reproductive needs, and habitat

preferences (Day, 2016; Rose et al., 2020; Wilke

et al., 2019).

Global results indicate that A. vittatus

is typically found in areas with temperatures

averaging between 24 and 26°C and precipi-

tation ranging from 42 to 101 mm. Species

of medical and veterinary importance, such

as A. aegypti and A. albopictus, which have a

tropical and subtropical distribution, exhibit

16 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

optimal development temperatures between 20

and 32 °C (Reinhold et al., 2018), while spe-

cies of the genus Culex thrive at optimal tem-

peratures between 24 and 30 °C (Ciota et al.,

2014). Regarding A. vittatus, a study assessing

the potential distribution of arbovirus vectors

in Morocco suggests that it may be found in

regions with a mean annual temperature rang-

ing from 20 to 30 °C (Abdelkrim et al., 2021).

However, the mean temperature for Europe

was found to be 15.4 °C. Nevertheless, our

simulations under current conditions estimate

suitable areas on this continent. Reproductive

success in colder zones may be attributed to the

diapause capacity observed in eggs and larvae

of species from genera such as Aedes, Anoph-

eles, Ochlerotatus, Orthopodomyia, among oth-

ers (Denlinger & Armbruster, 2014; Diniz et al.,

2017), as well as population growth during the

summer or until conditions become optimal for

development (Fonseca et al., 2015).

Based on occurrence records, A. vitta-

tus is primarily found in tropical forests and

tropical grasslands, followed by dry (desert)

and temperate habitats (Mediterranean forests,

temperate forests, and temperate grasslands).

At present, there are no studies that provide a

detailed description or specification of the hab-

itats where A. vittatus is globally distributed.

However, studies in Africa for A. vittatus and

other mosquito species have documented their

abundance in arid lands, savannas, forests, agri-

cultural soils, and villages in which it has been

dominant in microhabitats, like puddles, fruits,

rock pools and tree holes. In the absence of nat-

ural breeding sites, the vector can reproduce in

artificial containers such as plastic bottles, cups,

used cans, broken clay pots, and so on (Ali et

al., 2014; Diallo et al., 2012a; Obi et al., 2022).

Models obtained under current conditions

estimate that A. vittatus is primarily distributed

in the tropical region (up to 3 900 m for the

African hypothesis and 4 900 m for the Asian

hypothesis) and in some subtropical zones (up

to 2 900 m for the African hypothesis and 3 900

m for the Asian hypothesis). Currently, it is sug-

gested that A. vittatus is mostly distributed in

the tropical region with suitable areas of lower

probability in subtropical regions (Abdelkrim et

al., 2022). Invasive species at a global scale, such

as A. aegypti (African) and A. albopictus (Asian)

have distributions in both regions. Specifically,

A. aegypti exhibits a wider invasion in tropical

regions compared to subtropical regions. In

contrast, A. albopictus has a broad invasion in

both tropical and subtropical regions, and it is

estimated to have established in areas including

Europe, Iceland, Greenland, the United States,

and Japan (Kamal et al., 2018; Kraemer et al.,

2015). According to our estimations, A. vittatus

could potentially exhibit a pattern similar to

A. albopictus. Regardless of its native area, this

mosquito has the potential to invade tropical

and subtropical zones with altitudes exceeding

2 000 m. Furthermore, under future condi-

tions, it is estimated that the invasion of both

A. aegypti and A. albopictus will increase. In

contrast, for A. vittatus, its range of invasion is

expected to decrease in high emission scenarios

(RCP 8.5) in areas such as Spain, Portugal,

Australia, Morocco, and Algeria (Abdelkrim et

al., 2022). However, our estimations indicate a

decrease in Australia for the period 2081-2100

in both hypotheses. Additionally, reduced inva-

sion is also projected in other regions of Africa,

Asia, America, and Oceania under scenario

SSP585. In contrast, our results for the period

2081-2100, under the Asian hypothesis, sug-

gest that A. vittatus could be present in areas of

Greenland and Iceland, similar to the estimates

for A. albopictus in 2050 (Kamal et al., 2018).

In both hypotheses, invasion is projected

to occur in tropical and subtropical regions.

In the case of the native African hypothesis, it

suggests a higher level of invasion toward the

European continent and North America. Con-

versely, for the Asian hypothesis, the distribu-

tion and invasion rates at these sites are lower.

In the future, under the African hypothesis,

invasion is projected to extend further into the

tropical and subtropical zones of all continents,

including Asia. Conversely, under the Asian

hypothesis, the extension into Europe remains

limited, with a notable increase in Asia, par-

ticularly towards cold zones such as Greenland

and Antarctica. Global transport networks,

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e54166, enero-diciembre 2024 (Publicado May. 21, 2024)

climate change, and increased urbanization

have facilitated the spread of invasive species.

Additionally, their ability to colonize new ter-

ritories leads to economic, environmental, and

public health damage due to various human

activities that favor their proliferation (Schaff-

ner et al., 2013; Weaver & Reisen, 2010). As a

result, invasive species such as A. aegypti and

A. albopictus have expanded their distribution

range (Kraemer et al., 2019). In this case, sea

lanes play a crucial role in the distribution and

invasion of vectors at continental scales (Louni-

bos, 2002; Service & Place, 1997) as they enable

access to new sites through seaports (Wilke et

al., 2022). Consequently, A. vittatus, which had

previously been documented only in Africa,

Asia, and occasionally in Europe, has recently

been reported in the Americas. Estimations

suggest a future cosmopolitan behavior for this

vector, with the potential to be found in conti-

nents where it has not been previously recorded

and the possibility of colonizing colder cli-

mates, including suitable areas in Greenland,

Iceland, and Antarctica.

In conclusion, A. vittatus exhibits a tropical

and subtropical distribution, inhabiting rural,

urban, and peri-urban areas at altitudes ranging

from sea level to 2 500 m.a.s.l., with tempera-

tures between 15 and 30 °C, in various land-

scapes such as forests, grasslands, deserts, and

lakes. The mosquito has the potential to invade

new areas in Europe, America, and Oceania.

Climate change is expected to drive invasions

towards colder subtropical zones in coun-

tries across Asia, Europe, and North America,

regardless of its native range. In a period of

around 80 years (2081-2100), invasion is sug-

gested toward zones of Greenland and the Ant-

arctic with the hypothesis that A. vittatus being

native of Asia and reaching altitudes up to 5 000

m in China under the three scenarios. There-

fore, given A. vittatus’ potential for invasion

and ecological adaptability, vector control and

epidemiological disease management programs

should include monitoring for the presence of

this vector in the locations described in this

study. Likewise, it is important that in the areas

where the vector has been recorded, studies

are conducted to understand its population

dynamics, feeding and reproductive behavior,

life cycles, and preferred habitats. Similarly,

given the recent knowledge of its invasion in

the Americas, studies aimed at understanding

its population dynamics and potential spread to

new locations are recommended.

Ethical statement: the authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

See supplementary material

a27v72n1-MS1

ACKNOWLEDGMENTS

The authors would like to express their

gratitude to the Vice-rectory of Research at

Universidad del Quindío for funding the trans-

lation of the manuscript. Additionally, special

thanks are extended to Chrystian Camilo Sosa

Arango, MSc., for their invaluable support in

constructing the models.

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