Aedes albopictus - Factsheet for experts

Hazard associated with mosquito species

Current issues

Invasive species/global dispersion

Aedes albopictus has undergone a dramatic global expansion facilitated by human activities, in particular the movement of used tyres and ‘lucky bamboo’ [1]. Together with passive transit via public and private transport, this has resulted in a widespread global distribution of Ae. albopictus. It is now listed as one of the top 100 invasive species by the Invasive Species Specialist Group [2].

Ecological plasticity

The success of the invasion of Ae. albopictus is due to a number of factors including: its ecological plasticity, strong competitive aptitude, globalization i.e. increase of trade and travel, lack of surveillance, and lack of efficient control [1]. Climate change predictions suggest Ae. albopictus will continue to be a successful invasive species that will spread beyond its current geographical boundaries [3-5]. This mosquito is already showing signs of adaptation to colder climates [1,6] which may result in disease transmission in new areas.

Biting and disease risk

Aedes albopictus feeds on a wide range of hosts. It is also known to be a significant biting nuisance, with the potential to become a serious health threat as a bridge vector of zoonotic pathogens to humans [7]. This mosquito species is a known vector of chikungunya virus, dengue virus and dirofilariasis. A number of other viruses affecting human health have also been isolated from field-collected Ae. albopictus in different countries. Moreover, its recent involvement in the localised transmission of chikungunya virus in Italy [8] and France [9,10] and dengue virus in France [11-13] and Croatia [14] highlights the importance of monitoring this invasive species.

Geographical distribution

Aedes albopictus has been reported in the following areas: [1,3,6,7,15-42].

• Europe: Albania, Austria (not established to date), Belgium (not established to date), Bosnia & Herzegovina, Bulgaria, Croatia, Czech Republic (not established to date), France (including Corsica), Georgia, Germany, Greece, Hungary, Italy (including Sardinia, Sicily, Lampedusa, and other islands), Malta, Monaco, Montenegro, the Netherlands (not established to date), Romania, Russia, San Marino, Serbia (not established to date), Slovakia (not established to date), Slovenia, Spain, Switzerland, Turkey and Vatican City
• Middle East: Israel, Lebanon, Saudi Arabia (to be confirmed), Syria, Yemen (to be confirmed)
• Asia & Australasia: Australia (established only in the Torres Strait, the region that separates mainland Australia from Papua New Guinea), Japan, New Zealand (not established), numerous Pacific Ocean and Indian Ocean islands, southern Asia
• North, Central America & Caribbean: Barbados (not established), Belize, Cayman Islands, Costa Rica, Cuba, Dominican Republic, El Salvador, Guatemala, Haiti, Honduras, Mexico, Nicaragua, Panama, Trinidad (not established), USA
• South America: Argentina, Bolivia (not confirmed), Brazil, Colombia, Paraguay, Uruguay, Venezuela
• Africa: Algeria, Cameroon, Central African Republic, Equatorial Guinea, Gabon, Madagascar, Nigeria, Republic of Congo, South Africa (not established)

The current known distribution of Ae. albopictus in Europe in displayed on the vector maps

Brief history of spread and European distribution

Pathways

Having originated in tropical forests of South-East Asia, Ae. albopictus has spread globally. This geographical spread has mostly occurred during the past three decades [1] via passive transport of eggs in used tyres or lucky bamboo, the latter being the route of importation into Belgium, the Netherlands and California [16,43,44]. Public or private transport from heavily-infested areas has also resulted in the passive transportation of Ae. albopictus into new areas. Passive transport from heavily infested areas via ground vehicles is believed to be the route of introduction of Ae. albopictus into southern France, Germany, the Balkans, the Czech Republic, Spain and Switzerland [19,23,30,45].

Timeline of initial movements

Aedes albopictus was first reported in Europe in 1979 in Albania [46]. In 1985 it was reported in Texas, USA and has since spread northward and eastward, having now been reported in at least 32 US states including Hawaii [38]. This expansion was facilitated by the movement of used tyres along the interstate highways [18]. In Latin America it was first reported in Brazil in 1986 and later in Mexico in 1988 [19]. In Africa, it was first detected in 1990 in South Africa but establishment was only reported in 2000 from Cameroon [47,48].

Initial importation and spread in Europe

The first record of importation to Europe was in Albania in 1979 but it was suspected to be present from 1976. Although Ae. albopictus became established in Albania, there were no reports in any other European country until 1990, when it was found in Italy [49]. Since its importation into Italy through Genoa [23], Ae. albopictus has now become established in most areas of the country <600m above sea level and is abundant in many urban areas [3,50]. During the first 10 years of colonisation in the country, Ae. albopictus spread throughout 22 provinces, mainly in the north east of the country [51]. Italy is now the most heavily-infested country in Europe, with the highest incidence in the Veneto and Friuli-Venezia-Giulia regions, large parts of Lombardia and Emilia-Romagna and coastal areas of central Italy [3]. The mosquito was then reported in France in 1999 and Belgium in 2000. These initial importations were subsequently eradicated or died out, but spread has occurred to a number of countries in Europe since 2000, including southern France.

Having become established in Albania, Italy and on the Cote d’Azur in France, Ae. albopictus is also known to be spreading in Greece, Spain and the Balkan countries [23]. Aedes albopictus has also been reported in Ticino in Switzerland since 2003, suggesting sporadic introductions from Italy [52]. In 2004, it was reported near Barcelona in Spain, with some spread along the Mediterranean coast [19]. Furthermore, it has been repeatedly found in the Netherlands (2005, 2006 and 2007) at the premises of companies importing bamboo [43,53] and in Malta [26]. The Dutch populations, imported with lucky bamboo, have not established outside greenhouses, suggesting that they are tropical strains. Besides, populations are also imported from USA via used tyre trade, and control measures have so far avoided their establishment [54]. Aedes albopictus has been trapped on a number of occasions along motorways in southern Germany, suggesting introduction by vehicles from southern Europe [32,45,55]. In 2014, all developmental stages were found over extended periods of time in southern Germany indicating local reproduction [56].

Further specimens were collected at parking lots along motorways and at some other places in Austria [31], Czech Republic [30], and Slovakia [33], but subsequent surveys remained negative [38]. Finally, populations have been found established in Slovenia in 2007 [57,58], in Bulgaria in 2011 [59], Russia in 2011 [60,61], Turkey in 2011 [35] and Romania in 2012 [40]. 

Possible future expansion 

The ability for imported populations to establish is currently dependent on the origin of the mosquito and its strain and it is not always clear whether introductions into Europe will result in established populations. In this context, it is suggested that Portugal, the eastern Adriatic Coast, eastern Turkey and the Caspian Sea coast of Russia are the most likely places for Ae. albopictus to establish itself in Europe [7]. Risk mapping projections suggest that further expansion of this species will occur in the Mediterranean basin towards the east and the west, as well as in the coastal areas of Greece, Turkey and the Balkan countries. Incorporation of climate change projections suggests that over time most of Europe will become more suitable for Ae. albopictus establishment [3,62,63]. Especially Western Europe (Belgium, France, Luxembourg and the Netherlands) will provide favourable climatic conditions within the next decades. Climatic conditions will continue to be suitable in southern France as well as in most parts of Italy and Mediterranean coastal regions in south-eastern Europe [63]. It is predicted that future climate trends will increase the risk of establishment in northern Europe, e.g. parts of Germany and the southernmost parts of the UK, due to wetter and warmer conditions, and slightly decrease the risk across southern Europe because of hotter and drier summers [3,62,63]. Land use changes, particularly urbanisation, may continue to increase the competitive advantage of Ae. albopictus over resident mosquitoes through its exploitation of artificial container habitats; further aiding establishment in new areas [64]. Winter temperatures and mean annual temperatures appear to be the most significant limiting factors of Ae. albopictus expansion in Europe [65].

Entomology

• SPECIES NAME/CLASSIFICATION: Aedes (Stegomyia) albopictus (Skuse) [66]
• COMMON NAME: Asian tiger mosquito, Forest day mosquito
• SYNONYMS AND OTHER NAME IN USE: Stegomyia albopicta (sensu Reinert et al. [67]) 

Morphological characters and similar species

Aedes albopictus adults are relatively small and show a black and white pattern due to the presence of white/silver scale patches against a black background on the legs and other parts of the body. Some indigenous mosquitoes also show such contrasts but these are less obvious (more brownish and yellowish). Aedes albopictus can, however, be confused with other invasive (Ae. aegypti, Ae. japonicus) or indigenous species (Ae. cretinus, restricted to Cyprus, Greece and Turkey), and the diagnostic character is the presence of a median silver-scale line against a black background on the scutum (dorsal part of the thorax). The differentiation with Ae. cretinus needs a detailed check of scale patches on the thorax.

Life history

Diapausing tendencies

Tropical and subtropical populations are active throughout the year with no diapausing phase [26]. Temperate populations are affected by seasonal temperature and photoperiodicity and, in response to these factors, can overwinter by producing eggs that undergo a winter diapause [68]. Eggs, laid during late summer or early autumn when daylight hours are reducing, enter facultative diapause, and hatching suppression occurs which is usually sufficient to outlast winter [68]. The species’ ability to induce photoperiodic egg diapause allows it to overwinter in temperate regions, which assists its establishment in more northern latitudes in Asia, North America and Europe. Diapausing eggs of European Ae. albopictus have been shown to be able to survive a cold spell of -10oC, whereas eggs of tropical Ae. albopictus could only survive -2oC. In addition, the hatching success and cold tolerance of European Ae. albopictus eggs were increased in diapausing eggs when compared to non-diapausing eggs [69]. Aedes albopictus populations in Italy are showing signs of cold-acclimation as adults and are thus remaining active throughout winter [70]. Some populations in North America are likely to be exposed to mean temperatures of -5˚C and will overwinter if females have deposited eggs in containers that are not exposed to these temperatures for prolonged periods ‒ e.g. artificial containers in peridomestic areas [64].

General life history

The drought-resistant eggs are laid above the water line. Larval/pupal development takes three to eight weeks and is continuous throughout the year in European southernmost regions (Malta [26]). Adult females can survive over three weeks [70]. They have been reported to overwinter in Rome [70] and even to lay eggs during winter time in Spain [71].

Seasonal abundance

Seasonal abundance is dependent on temperature and the availability of food and water in a particular geographical area. Higher temperatures speed up larval development, increasing the number of adult populations, the autumnal development of immatures and consequently the rates of egg overwintering [68]. A study in northern Italy showed an increased abundance of adult females during the period May-September, peaking in late July [72]. In Greece, Ae. albopictus is continuously active for over eight months of the year with the greatest abundance during summer and autumn, peaking in October. Oviposition takes place from mid-April to December, with the numbers of eggs highest from mid-July to the end of the autumn, and significantly increased during mild and rainy weather [73].

Voltinism (generations per season)

Multivoltine, 5-17 generations per year [26].

Host preferences

Aedes albopictus is an opportunistic feeder [74]. Blood hosts include humans, domestic and wild animals, reptiles, birds and amphibians [18]. Yet, laboratory studies and blood meal analysis have shown a preference for human blood meals [1]. A study in Italy found a preference for mammals as opposed to birds and found human blood meals were more frequent in urban areas than rural sites, suggesting that host availability and abundance has a direct impact on the feeding activities of Ae. albopictus [50].

Aquatic and terrestrial habitats

Aedes albopictus has the ability to breed in natural and artificial habitats, some of which include tyres, barrels, rainwater gulley catch basins and drinking troughs [26]. Natural habitats consist in phytotelms (water bodies held by terrestrial plants e.g. tree holes) and rock pools [75]. They are not known to breed in brackish or salt water [24]. In general, albeit in Europe, they have a preference for urban and suburban habitats [76]. Aedes albopictus is said to be superior in competing for food resources with Ae. triseriatus and Ae. japonicus [64].

Biting and resting habits

Aedes albopictus is currently considered a serious biting nuisance for humans in Italy [23,77], southern France [78] and Spain where it is significantly reducing the quality of life in infested areas [19]. Adult females bite aggressively, usually during the day and preferably outdoors. However, there are reports that Ae. albopictus is becoming partially endophilic [77], and is found to be biting indoors [79]. During a study in Rome, blood-fed females were mainly found indoors, indicating that local mosquito populations could spend time resting indoors after a blood meal [50]. Another study on Penang Island in Malaysia reported observations of Ae. albopictus females developing indoors within containers. Such containers included flower vases, empty paint cans and sinks. Most stages of larval populations were present over a five-month period, suggesting that this species may have adapted to indoor environments [80]. A laboratory study found that Ae. albopictus could survive for long periods indoors by obtaining sugars from lucky bamboo and other ornamental plants [81]. The mosquitoes’ survival time was long enough to complete a gonotrophic cycle, and to allow development of transmissible arboviruses within the vector [81].

Environmental thresholds/constraints/development criteria

Establishment thresholds

The extent to which Ae. albopictus has established itself in new geographic locations is thought to correspond to various climatic thresholds: a mean winter temperature of >0oC to permit egg overwintering, a mean annual temperature of >11oC required for adult survival and activity, and at least 500mm of annual rainfall; a pre-requisite for the maintenance of aquatic habitats. However, rainfall needs to be sufficient during summer months to maintain such sites [68,82]. Conversely, periods of high precipitation reduce short-term abundance of host-seeking females [72]. A summer temperature of 25‒30oC is required for optimum development [83]. However, there are reports of populations establishing in areas with lower mean temperatures (5‒28.5°C) and lower rainfall (290mm annually) than previously suspected [7,84].

Diapausing and reactivation cues

The length of the reproductive season is regulated by the increasing temperatures in spring and the onset of egg diapause in autumn. The critical photoperiodic threshold varies between geographical locations. In general, the production of diapausing eggs occurs below 13‒14 hours of daylight, however in some locations this threshold occurs at 11‒12 hours [68].

Hatching of diapausing eggs in spring is related to changes in photoperiod (i.e. length of day), food availability, temperature, and water availability. Furthermore, this mosquito may not survive through winter if environmental temperatures and humidity are not maintained above set thresholds, or if the diapause period exceeds six months [68].

There is generally little adult activity below 9oC, but adults do seek warmer microclimates indoors [72]. In parts of Italy, adult activity continues throughout winter [70].

Dispersal range

Flight range (and hence dispersal on the wing) is limited to ~200m [74]. The main dispersal route for this mosquito is via transport and movement of container habitat goods.

Epidemiology and transmission of pathogens

Known vector status

During the 2006-2007 chikungunya outbreak in Italy, the status of Ae. albopictus as vector of the chikungunya virus was clearly demonstrated [1]. This mosquito is also known to be able to transmit dengue virus [85,86] and dirofilarial worms [19,87]. All four dengue virus serotypes have been isolated from Ae. albopictus [88]. Infection studies of l Ae. albopictus suggest a possible contribution to Zika virus outbreaks [89,90].

Aedes albopictus is considered to be a competent vector experimentally of at least 22 other arboviruses including yellow fever virus, Rift Valley fever virus, Japanese encephalitis virus, West Nile virus and Sindbis virus, all of which are relevant to Europe. Potosi virus, Cache Valley virus, La Crosse virus, Eastern equine encephalitis virus, Mayaro virus, Ross River virus, Western equine encephalitis virus, Venezuelan equine encephalitis virus, Oropouche virus, Jamestown Canyon virus, San Angelo virus and Trivittatus virus are other arboviruses that Ae. albopictus can transmit experimentally [38,91].

A number of these viruses have also been isolated from field-collected Ae. albopictus in different countries and laboratory transmission of such viruses by Ae. albopictus has been demonstrated [1]. These include Eastern equine encephalitis virus [92,93], La Crosse virus [94,95], Venezuelan equine encephalitis virus [96,97], West Nile virus [72,98,99] and Japanese encephalitis virus [1]. Usutu virus has been isolated from Ae. albopictus in Italy, but it is unknown whether the mosquito can transmit this pathogen [100]. Field isolation and experimental infection studies alone do not prove that this mosquito species is involved in the transmission of such viruses, but the mosquito’s biting habits, increasing global distribution and recent involvement in a chikungunya virus outbreak highlight the significance of Ae. albopictus to public health.

A high prevalence of the insect-infective Aedes flavivirus has been detected in Ae. albopictus in Italy and it has been suggested that its presence in these mosquitoes could influence the transmission dynamics of other human-pathogenic flaviviruses, such as West Nile virus and Usutu virus [101].

Not only does Ae. albopictus represent a disease risk but it can also cause a considerable amount of nuisance biting in areas where it is well-established, reducing the quality of life of individuals affected [102]. Prevalence of Ae. albopictus has also been linked to a reduction in children’s outdoor physical activity time, a factor contributing to childhood obesity [103].

Chikungunya

Aedes albopictus mosquitoes are able to transmit chikungunya virus within two days of ingesting a viraemic blood meal [104]. Some experts suggest that transovarial transmission is enough to maintain viral cycles but others disagree [23]. No evidence of transovarial transmission was found during an entomological investigation into the 2007 chikungunya outbreak in Italy [105] but the virus was detected in field-caught male Ae. albopictus following an outbreak in Thailand [106].

Chikungunya was first reported in Europe in 2007 following epidemics in the Indian Ocean (2005‒2007), which caused millions of cases and significant morbidity and burden on health resources. This was the first known local transmission of chikungunya in Europe and occurred in Emilia-Romagna province in Italy. An infected traveller returned home from India, spreading the disease to localised populations of Ae. albopictus mosquitoes. Following entomological investigations during the outbreak females of Ae. albopictus were found to be PCR positive and the virus was successfully isolated [105]. The adaptation of the virus to this new vector host (in addition to its principle vector Ae. aegypti) has resulted in improved virus replication and transmission efficiency of the virus by Ae. albopictus [5,107] [104]. Autochthonous chikungunya fever cases occurred in south eastern France in 2010 and 2014 [9,10]. Tilston et al considers that, based on temperature, the southern European countries are most at risk of chikungunya virus transmission [108].

Dengue

Although generally considered a secondary vector of dengue to Ae. aegypti, Ae. albopictus has been associated with dengue virus transmission and this has been acknowledged since the mid-nineteenth century [1]. It was implicated as the vector responsible for outbreaks in Hawaii [85] Reunion Island and Mauritius [86,109] . It has also been associated with dengue virus transmission in China, Japan and Seychelles [88]. Dengue virus is transmitted transovarially so emergence of adults from imported infected eggs could lead to further spread of the disease [24]. Dengue virus can also be transmitted venereally in mosquitoes [88].

Autochthonous cases of dengue were reported in France during September 2010 [11] followed by others in Croatia at around the same time [14]. Further cases, linked to Ae. albopictus, were reported from France in 2013, 2014 and 2015 [12,13,110]. Although modelling predicts that most of Europe is currently unsuitable for dengue transmission, areas combining high human population density with suitable day- and night-time land surface temperatures are still at greater risk [4,111]. However, competent mosquito vectors must be present for transmission to occur. Climatically, areas predicted to be at risk from dengue include northern Italy, parts of Austria, Slovenia and Croatia, and west of the Alps in France [111]. The risk of transmission to humans is considered to be higher where there is a presence of Ae. aegypti than in areas with Ae. albopictus. This point is exemplified by the outbreak of dengue on Madeira associated with Ae. aegypti [112].

Zika

Aedes albopictus is considered a potential vector of Zika virus. Vector competence studies of local Ae. albopictus in Singapore with the African lineage of Zika virus showed the potential of this mosquito to transmit Zika virus [89]. Recent studies using different Ae. albopictus populations from the Americas and Europe revealed that this mosquito is susceptible to Zika virus infection, that the virus is disseminated and can reach the salivary glands but not very efficiently; Ae. albopictus has a lower vector competence compared to Aedes aegypti  [113] [114] [115]. The species has been found infected in wild caught mosquitoes [90].

Dirofilariasis

Aedes albopictus has a role in the transmission of Dirofilaria in Asia, North America and Europe [1]. Dirofilaria (filarial nematodes D. immitis and D. repens) is a parasite transmitted primarily between dogs (or other canids which act as reservoir hosts) and mosquitoes, but which can also affect humans. Recent evidence has shown transmission of the parasite by Italian Ae. albopictus populations [116-118], coupled with an increase in prevalence of human dirofilariasis in Italy [87].

Human infections are increasing in Europe and although it is unusual for the parasite to develop into the adult stage in humans, at least three cases of microfilaraemic zoonotic infections have been reported in Europe [77,119].

Factors driving/impacting on transmission cycles

A growth in dengue cases worldwide, increasing global travel and established populations of Ae. albopictus may have been the cause of the dengue outbreak in Mauritius in June 2009 [86]. The movement of viraemic hosts can result in outbreaks of chikungunya virus in non-endemic areas. Climate change could increase the distribution of Ae. albopictus beyond its current boundaries which could enhance the transmission potential of chikungunya virus and dengue virus in temperate regions [5,77,120]. Aedes albopictus mosquitoes tend to feed on multiple hosts which also increases the risk of zoonotic disease transmission [1]. The return of viraemic travellers from disease-epidemic areas to temperate regions has resulted in (and will potentially continue to result in) local mosquito populations sustaining disease transmission [83]. Therefore, the presence of Ae. albopictus in Europe and the increasing number of overseas travellers may increase the risk of dengue and chikungunya outbreaks in Europe [121].

Public health (control/interventions)

Vector surveillance

Methods for surveying Ae. albopictus are addressed in the ‘ECDC Guidelines for the surveillance of invasive mosquitoes in Europe [122].

ECDC and EFSA fund European-wide monitoring and mapping activities for invasive mosquito species and potential mosquito vectors (VectorNet).

Species specific control methods

Source reduction and adult control

Control of Ae. albopictus is based on the reduction of larval development sites. Mosquito fogging and larviciding (insecticides targeting the mosquito larvae) were techniques used during an outbreak of dengue in Mauritius in June 2009 [86]. In 2006, use of insecticides in greenhouses that had been recently colonised by Ae. albopictus in the Netherlands may have contributed to the decline in numbers caught the following year [43]. Permethrin, Bacillus thuringiensis israeliensis ser. H14 and diflubenzuron (an insect growth regulator) were used to treat stagnant water after the detection of Ae. albopictus in Switzerland in 2003 [52]. Although resistance to insecticides is not currently a problem, it has been detected in a population in Thailand [123] and more recently in populations in La Reunion [124] and Malaysia [123,125]. In Pakistan, field-collected Ae. albopictus displayed moderate-high resistance to many agricultural insecticides, including pyrethroids [126].

Control of this species in newly-established areas has been difficult (e.g. USA, France and Italy) [1]. Although prevention of invasion was achieved after its first introduction into France in 1999, subsequent introductions (most likely by vehicles from Italy) have resulted in Ae. albopictus becoming a pest problem in southern France and Corsica. A study in Catalonia, Spain demonstrated the use of multiple intervention strategies (source reduction, larvicide and adulticide treatments and the cleaning of uncontrolled landfills) as successful in curbing an established population of Ae. albopictus (produced a marked reduction in egg numbers). The authors concluded that citizen cooperation was an essential component for successfully implementing these interventions [102].

Implementation of an integrated control strategy against invasive mosquito species should take into account the target species, its ecology and the public health concern, i.e. nuisance or disease transmission. As a general rule, an integrated Invasive Mosquito Species control strategy requires the coordinated involvement of local authorities, private partners, organised society and communities [127].

Decreasing human-vector contact and the use of public health material have been widely used in endemic areas in Europe, such as Italy. The use of irradiated or genetically modified mosquitoes which are still under development are methods that may be used in the future to complement conventional methods. Additional control methods which may be applied in the future include Wolbachia infection to block transmission of dengue virus and chikungunya virus, and the introduction of natural predators [34]. Personal protective measures to reduce the risk of mosquito bites include the use of mosquito bed nets (preferably insecticide-treated nets), sleeping or resting in screened or air-conditioned rooms, the wearing of clothes that cover most of the body, and the use of mosquito repellent in accordance with the instructions indicated on the product label.

Existing public health awareness and education materials

• ECDC also provides information on dengue, chikungunya, yellow fever and Zika virus.
• The ECDC provides updated vector distribution maps and step-by-step web guidelines for the surveillance of invasive mosquitoes.
• The CDC provides advice for travellers on protection against mosquitoes, ticks and other arthropods.
• The National travel health network and centre provides information on how to avoid insect bites (including mosquito bites).
• The Regional Committee for Europe has endorsed at his 63rd session, September 2013, a ‘Regional framework for surveillance and control of invasive mosquito vectors and re-emerging vector-borne diseases 2014–2020’ [report] [resolution

Key areas of uncertainty 

Although it is not clear how significant Ae. albopictus will be in disease transmission across Europe, the ability of Ae. albopictus to adapt to new environments, its predicted spread and establishment in Europe and its confirmed involvement in disease transmission cycles makes the surveillance and control of this species hugely important. 

References

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26.          Gatt P, Deeming JC, Schaffner F. First records of Aedes (Stegomyia) albopictus (Skuse) (Diptera: Culicidae) in Malta. Eu Mosq Bull. 2009;27 56-64.
27.          Carrieri M, Albieri A, Angelini P, Baldacchini F, Venturelli C, Zeo SM, et al. Surveillance of the chikungunya vector Aedes albopictus (Skuse) in Emilia-Romagna (northern Italy): organizational and technical aspects of a large scale monitoring system. J Vector Ecol. 2011 Jun;36(1):108-16.
28.          Izri A, Bitam I, Charrel RN. First entomological documentation of Aedes (Stegomyia) albopictus (Skuse, 1894) in Algeria. Clin Microbiol Infect. 2011;17(7):1116-8.
29.          Fernandez Mdel C, Jean YS, Callaba CA, Lopez LS. The first report of Aedes (Stegomyia) albopictus in Haiti. Memórias do Instituto Oswaldo Cruz. 2012 Mar;107(2):279-81.
30.          Sebesta O, Rudolf I, Betasova L, Pesko J, Hubalek Z. An invasive mosquito species Aedes albopictus found in the Czech Republic, 2012. Euro Surveill. 2012;17(43):20301.
31.          Seidel B, Duh D, Nowotny M, Allerberger F. Erstnachweis der Stechmücken Aedes (Ochlerotatus) japonicus japonicus (Theobald, 1901) in Österreich und Slowenien in 2011 und für Aedes (Stegomyia) albopictus (Skuse, 1895) in Österreich 2012 (Diptera: Culicidae). Entomologische Zeitschrift. 2012;122(5):223-6.
32.          Becker N, Geier M, Balczun C, Bradersen U, Huber K, Kiel E, et al. Repeated introduction of Aedes albopictus into Germany, July to October 2012. Parasitol Res. 2013 Apr;112(4):1787-90.
33.          Bockova E, Kocisova A, Letkova V. First record of Aedes albopictus in Slovakia. Acta Parasitologica 2013;58(4):603-6.
34.          Bonizzoni M, Gasperi G, Chen X, James AA. The invasive mosquito species Aedes albopictus: current knowledge and future perspectives. Trends Parasitol. 2013 Sep;29(9):460-8.
35.          Oter K, Gunay F, Tuzer E, Linton YM, Bellini R, Alten B. First record of Stegomyia albopicta in Turkey determined by active ovitrap surveillance and DNA barcoding. Vector Borne Zoonotic Dis. 2013 Oct;13(10):753-61.
36.          Carvalho RG, Lourenco-de-Oliveira R, Braga IA. Updating the geographical distribution and frequency of Aedes albopictus in Brazil with remarks regarding its range in the Americas. Memórias do Instituto Oswaldo Cruz. 2014 Sep;109(6):787-96.
37.          Adeleke MA, Sam-Wobo SO, Garza-Hernandez JA, Oluwole AS, Mafiana CF, Reyes-Villanueva F, et al. Twenty-three years after the first record of Aedes albopictus in Nigeria: Its current distribution and potential epidemiological implications. African Entomology 2015;23(2):348-3.
38.          Medlock JM, Hansford KM, Versteirt V, Cull B, Kampen H, Fontenille D, et al. An entomological review of invasive mosquitoes in Europe. Bulletin of Entomological Research. 2015 Dec;105(6):637-63.
39.          Ngoagouni C, Kamgang B, Nakoune E, Paupy C, Kazanji M. Invasion of Aedes albopictus (Diptera: Culicidae) into central Africa: what consequences for emerging diseases? Parasit Vectors. 2015;8:191.
40.          Prioteasa LF, Dinu S, Falcuta E, Ceianu CS. Established Population of the Invasive Mosquito Species Aedes albopictus in Romania, 2012-14. J Am Mosq Control Assoc. 2015 Jun;31(2):177-81.
41.          Akiner MM, Demirci B, Babuadze G, Robert V, Schaffner F. Spread of the Invasive Mosquitoes Aedes aegypti and Aedes albopictus in the Black Sea Region Increases Risk of Chikungunya, Dengue, and Zika Outbreaks in Europe. PLoS Negl Trop Dis. 2016;10(4):e0004664.
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45.          Kampen H, Kronefeld M, Zielke D, Werner D. Further specimens of the Asian tiger mosquito Aedes albopictus (Diptera, Culicidae) trapped in southwest Germany. Parasitol Res. 2013 Feb;112(2):905-7.
46.          Adhami J, Reiter P. Introduction and establishment of Aedes (Stegomyia) albopictus skuse (Diptera: Culicidae) in Albania. J Am Mosq Control Assoc. 1998 Sep;14(3):340-3.
47.          Cornel AJ, Hunt RH. Aedes albopictus in Africa? First records of live specimens in imported tires in Cape Town. 1991 Mar;7(1):107-8.
48.          Fontenille D, Toto JC. Aedes (Stegomyia) albopictus (Skuse), a potential new Dengue vector in southern Cameroon. Emerg Infect Dis. 2001;7(6):1066-7.
49.          Sabatini A, Raineri V, Trovato G, Coluzzi M. [Aedes albopictus in Italy and possible diffusion of the species into the Mediterranean area]. Parassitologia. 1990 Dec;32(3):301-4.
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52.          Wymann MN, Flacio E, Radczuweit S, Patocchi N, Luthy P. Asian tiger mosquito (Aedes albopictus) - a threat for Switzerland? Euro Surveill. 2008 Mar 6;13(10):8058.
53.          Scholte EJ, Dijkstra E, Ruijs H, Jacobs F, Takken W, Hofhuis A, et al. The Asian tiger mosquito in the Netherlands: should we worry? . Proceed Section Exp Appl Entomol. 2007;18 131-6.
54.          Scholte E, Den Hartog W, Dik M, Schoelitsz B, Brooks M, Schaffner F, et al. Introduction and control of three invasive mosquito species in the Netherlands, July-October 2010. Euro Surveill. 2010;15(45):19710.
55.          Werner D, Kronefeld M, Schaffner F, Kampen H. Two invasive mosquito species, Aedes albopictus and Aedes japonicus japonicus, trapped in south-west Germany, July to August 2011. Euro Surveill. 2012;17(4):20067.
56.          Werner D, Kampen H. Aedes albopictus breeding in southern Germany, 2014. Parasitol Res. 2015 Mar;114(3):831-4.
57.          Kalan K, Kostanjšek R, Merdić E, Trilar T. A survey of Aedes albopictus (Diptera: Culicidae) distribution in Slovenia in 2007 and 2010. Natura Sloveniae. 2011;13(1):39-50.
58.          Kalan K, Buzan VE, Ivović V. Distribution of two invasive mosquito species in Slovenia in 2013. Parasit Vectors. 2014;7(Suppl 1)(P9).
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24.          Buhagiar JA. A second record of Aedes (Stegomyia) albopictus (Diptera: Culicidae) in Malta. Eu Mosq Bull. 2009;27:65-7.
25.          Diallo M, Laganier R, Nangouma A. First record of Ae. albopictus (Skuse 1894), in Central African Republic. Trop Med Int Health. 2010;15(10):1185-9.
26.          Gatt P, Deeming JC, Schaffner F. First records of Aedes (Stegomyia) albopictus (Skuse) (Diptera: Culicidae) in Malta. Eu Mosq Bull. 2009;27 56-64.
27.          Carrieri M, Albieri A, Angelini P, Baldacchini F, Venturelli C, Zeo SM, et al. Surveillance of the chikungunya vector Aedes albopictus (Skuse) in Emilia-Romagna (northern Italy): organizational and technical aspects of a large scale monitoring system. J Vector Ecol. 2011 Jun;36(1):108-16.
28.          Izri A, Bitam I, Charrel RN. First entomological documentation of Aedes (Stegomyia) albopictus (Skuse, 1894) in Algeria. Clin Microbiol Infect. 2011;17(7):1116-8.
29.          Fernandez Mdel C, Jean YS, Callaba CA, Lopez LS. The first report of Aedes (Stegomyia) albopictus in Haiti. Memórias do Instituto Oswaldo Cruz. 2012 Mar;107(2):279-81.
30.          Sebesta O, Rudolf I, Betasova L, Pesko J, Hubalek Z. An invasive mosquito species Aedes albopictus found in the Czech Republic, 2012. Euro Surveill. 2012;17(43):20301.
31.          Seidel B, Duh D, Nowotny M, Allerberger F. Erstnachweis der Stechmücken Aedes (Ochlerotatus) japonicus japonicus (Theobald, 1901) in Österreich und Slowenien in 2011 und für Aedes (Stegomyia) albopictus (Skuse, 1895) in Österreich 2012 (Diptera: Culicidae). Entomologische Zeitschrift. 2012;122(5):223-6.
32.          Becker N, Geier M, Balczun C, Bradersen U, Huber K, Kiel E, et al. Repeated introduction of Aedes albopictus into Germany, July to October 2012. Parasitol Res. 2013 Apr;112(4):1787-90.
33.          Bockova E, Kocisova A, Letkova V. First record of Aedes albopictus in Slovakia. Acta Parasitologica 2013;58(4):603-6.
34.          Bonizzoni M, Gasperi G, Chen X, James AA. The invasive mosquito species Aedes albopictus: current knowledge and future perspectives. Trends Parasitol. 2013 Sep;29(9):460-8.
35.          Oter K, Gunay F, Tuzer E, Linton YM, Bellini R, Alten B. First record of Stegomyia albopicta in Turkey determined by active ovitrap surveillance and DNA barcoding. Vector Borne Zoonotic Dis. 2013 Oct;13(10):753-61.
36.          Carvalho RG, Lourenco-de-Oliveira R, Braga IA. Updating the geographical distribution and frequency of Aedes albopictus in Brazil with remarks regarding its range in the Americas. Memórias do Instituto Oswaldo Cruz. 2014 Sep;109(6):787-96.
37.          Adeleke MA, Sam-Wobo SO, Garza-Hernandez JA, Oluwole AS, Mafiana CF, Reyes-Villanueva F, et al. Twenty-three years after the first record of Aedes albopictus in Nigeria: Its current distribution and potential epidemiological implications. African Entomology 2015;23(2):348-3.
38.          Medlock JM, Hansford KM, Versteirt V, Cull B, Kampen H, Fontenille D, et al. An entomological review of invasive mosquitoes in Europe. Bulletin of Entomological Research. 2015 Dec;105(6):637-63.
39.          Ngoagouni C, Kamgang B, Nakoune E, Paupy C, Kazanji M. Invasion of Aedes albopictus (Diptera: Culicidae) into central Africa: what consequences for emerging diseases? Parasit Vectors. 2015;8:191.
40.          Prioteasa LF, Dinu S, Falcuta E, Ceianu CS. Established Population of the Invasive Mosquito Species Aedes albopictus in Romania, 2012-14. J Am Mosq Control Assoc. 2015 Jun;31(2):177-81.
41.          Akiner MM, Demirci B, Babuadze G, Robert V, Schaffner F. Spread of the Invasive Mosquitoes Aedes aegypti and Aedes albopictus in the Black Sea Region Increases Risk of Chikungunya, Dengue, and Zika Outbreaks in Europe. PLoS Negl Trop Dis. 2016;10(4):e0004664.
42.          Van den Hurk AF, Nicholson J, Beebe NW, Davis J, Muzari OM, Russell RC, et al. Ten years of the Tiger: Aedes albopictus presence in Australia since its discovery in the Torres Strait in 2005. One Health. 2016;2:19-24.
43.          Scholte EJ, Dijkstra E, Blok H, De Vries A, Takken W, Hofhuis A, et al. Accidental importation of the mosquito Aedes albopictus into the Netherlands: a survey of mosquito distribution and the presence of dengue virus. Med Vet Entomol. 2008 Dec;22(4):352-8.
44.          Demeulemeester J, Deblauwe I, De Witte J, Jansen F, Hendy A, Madder M. First interception of Aedes (Stegomyia) albopictus in Lucky bamboo shipments in Belgium. Journal of the European Mosquito Control Association 2014;32:14-6.
45.          Kampen H, Kronefeld M, Zielke D, Werner D. Further specimens of the Asian tiger mosquito Aedes albopictus (Diptera, Culicidae) trapped in southwest Germany. Parasitol Res. 2013 Feb;112(2):905-7.
46.          Adhami J, Reiter P. Introduction and establishment of Aedes (Stegomyia) albopictus skuse (Diptera: Culicidae) in Albania. J Am Mosq Control Assoc. 1998 Sep;14(3):340-3.
47.          Cornel AJ, Hunt RH. Aedes albopictus in Africa? First records of live specimens in imported tires in Cape Town. 1991 Mar;7(1):107-8.
48.          Fontenille D, Toto JC. Aedes (Stegomyia) albopictus (Skuse), a potential new Dengue vector in southern Cameroon. Emerg Infect Dis. 2001;7(6):1066-7.
49.          Sabatini A, Raineri V, Trovato G, Coluzzi M. [Aedes albopictus in Italy and possible diffusion of the species into the Mediterranean area]. Parassitologia. 1990 Dec;32(3):301-4.
50.          Valerio L, Marini F, Bongiorno G, Facchinelli L, Pombi M, Caputo B, et al. Host-feeding patterns of Aedes albopictus (Diptera: Culicidae) in urban and rural contexts within Rome province, Italy. Vector Borne Zoonotic Dis. 2010 Apr;10(3):291-4.
51.          Romi R, Di Luca M, Majori G. Current status of Aedes albopictus and Aedes atropalpus in Italy. J Am Mosq Control Assoc. 1999 Sep;15(3):425-7.
52.          Wymann MN, Flacio E, Radczuweit S, Patocchi N, Luthy P. Asian tiger mosquito (Aedes albopictus) - a threat for Switzerland? Euro Surveill. 2008 Mar 6;13(10):8058.
53.          Scholte EJ, Dijkstra E, Ruijs H, Jacobs F, Takken W, Hofhuis A, et al. The Asian tiger mosquito in the Netherlands: should we worry? . Proceed Section Exp Appl Entomol. 2007;18 131-6.
54.          Scholte E, Den Hartog W, Dik M, Schoelitsz B, Brooks M, Schaffner F, et al. Introduction and control of three invasive mosquito species in the Netherlands, July-October 2010. Euro Surveill. 2010;15(45):19710.
55.          Werner D, Kronefeld M, Schaffner F, Kampen H. Two invasive mosquito species, Aedes albopictus and Aedes japonicus japonicus, trapped in south-west Germany, July to August 2011. Euro Surveill. 2012;17(4):20067.
56.          Werner D, Kampen H. Aedes albopictus breeding in southern Germany, 2014. Parasitol Res. 2015 Mar;114(3):831-4.
57.          Kalan K, Kostanjšek R, Merdić E, Trilar T. A survey of Aedes albopictus (Diptera: Culicidae) distribution in Slovenia in 2007 and 2010. Natura Sloveniae. 2011;13(1):39-50.
58.          Kalan K, Buzan VE, Ivović V. Distribution of two invasive mosquito species in Slovenia in 2013. Parasit Vectors. 2014;7(Suppl 1)(P9).
59.          Mikov O, Nikolov G, Schaffner F, Mathis A, editors. First record and establishment of Aedes albopictus in Bulgaria. VBORNET-EMCA Joint Meeting ‘Invasive Mosquitoes and Public Health in the European Context’, ; 2013; Antwerp, Belgium, 28-29 November 2013.
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