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The Role of Culex pipiens L. (Diptera: Culicidae) in Virus Transmission in Europe

Victor A. Brugman, Luis M. Hernández-Triana, Jolyon M. Medlock and 3 more

International Journal of Environmental Research and Public Health · 2018 · Review · CC BY

doi:10.3390/ijerph15020389

Abstract

Over the past three decades, a range of mosquito-borne viruses that threaten public and veterinary health have emerged or re-emerged in Europe. Mosquito surveillance activities have highlighted the Culex pipiens species complex as being critical for the maintenance of a number of these viruses. This species complex contains morphologically similar forms that exhibit variation in phenotypes that can influence the probability of virus transmission. Critical amongst these is the choice of host on which to feed, with different forms showing different feeding preferences. This influences the ability of the mosquito to vector viruses and facilitate transmission of viruses to humans and domestic animals. Biases towards blood-feeding on avian or mammalian hosts have been demonstrated for different Cx. pipiens ecoforms and emerging evidence of hybrid populations across Europe adds another level of complexity to virus transmission. A range of molecular methods based on DNA have been developed to enable discrimination between morphologically indistinguishable forms, although this remains an active area of research. This review provides a comprehensive overview of developments in the understanding of the ecology, behaviour and genetics of Cx. pipiens in Europe, and how this influences arbovirus transmission.

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From PubMed Central, U.S. National Library of Medicine, as current on Oct 27, 2026. Paragraph numbers are ours, so an answer can point at one.

1. Introduction

Mosquitoes are responsible for the biological transmission of a wide diversity of arboviruses (arthropod-borne viruses) that cause diseases in humans, companion animals and livestock. Among the approximately 3500 mosquito species currently recognised worldwide, only a small number play a primary role in the transmission of arboviruses. The species that do fulfil this role tend to have adopted a degree of anthropophilic behaviour and occur at high abundance and in close proximity to susceptible hosts, primarily through exploitation of larval development sites created by humans.

In Europe, the recent emergence of mosquito-borne arboviruses has focused attention on identifying the species of mosquito that drive pathogen transmission. This phenomenon has occurred simultaneously with the incursion and establishment of several exotic and highly invasive species of mosquito associated with globalised trade. Several species of the Aedes genus have become established following importation and are a notable biting nuisance. Significantly, they change the epidemiological status of the region with respect to the occurrence of vector-borne disease. The first incursion into Europe of Aedes albopictus (Skuse, 1895) was reported in Albania in 1979, followed by Italy in 1990. The ability of this mosquito to exploit container habitats to breed in urban areas, produce diapausing eggs in temperate regions, and successfully expand through transportation in vehicles along highway systems, has facilitated its movement to more than 28 European countries and its establishment throughout large parts of the Mediterranean Basin, with a subsequent spread north. Similarly, since 2005, populations of Ae. aegypti (Linnaeus, 1762) have been reported on the Portuguese island of Madeira and are expanding in the Black Sea region; populations of Ae. japonicus (Theobald, 1901) have become widely established throughout Germany and have been reported from other countries such as Belgium and The Netherlands; and Ae. koreicus (Edwards, 1917) has been reported from Belgium and Italy. The involvement of Ae. albopictus in the local transmission of arboviruses previously considered to be exotic, such as chikungunya virus (CHIKV) in Italy, is now a major concern for European public health. This species may furthermore facilitate emergence and re-emergence of other viruses including dengue virus (DENV).

In contrast, invasive species of mosquito appear to have had only a limited impact on the transmission of arboviruses that have a longer history of circulation in Europe. The highest profile of these is West Nile virus (WNV), which has been present in this region for at least twenty years. In southern Europe WNV has been detected in the indigenous mosquito species Culex pipiens (L. 1758), which plays a primary role in transmission. This observation has been supported by laboratory studies that demonstrated susceptibility to infection in Cx. pipiens and isolated virus in saliva from fully susceptible individuals. Transmission rates of between 37% and 47% have been reported for populations in Italy where the virus is endemic, compared to 33% for populations tested from The Netherlands, suggesting that WNV could emerge in northern Europe.

There is no evidence that WNV has reached mosquito populations north of countries surrounding the Mediterranean Sea or south-east Europe, despite the presence of Cx. pipiens in many of these areas. However, the related flavivirus, Usutu virus (USUV), was detected in southern Europe around the same time as WNV, and has emerged in northern European countries including Germany, The Netherlands and Belgium. The principal vector for USUV is also Cx. pipiens and thus the reason for the absence of WNV in northern Europe is not fully understood, but among other factors may be related to the behaviour and distribution of different Cx. pipiens populations across Europe.

Previous reviews have considered the ecology of Cx. pipiens, current and future threats of mosquito-borne diseases across Europe and the influence of a changing climate on vector-borne disease. This review starts by presenting an overview of key arboviral threats to Europe, with focus on those for which Cx. pipiens is a vector. It then provides an updated overview of the literature relating to the taxonomy, ecology and behaviour of this important mosquito in Europe and examines future directions for research in these areas.

1.1. Viruses Associated with Transmission by Culex pipiens

At least ten arboviruses of medical and veterinary importance that are thought to be primarily transmitted by mosquitoes are currently circulating in Europe (Table 1). Culex pipiens has been shown to play a critical role in the transmission of three of these viruses. Conversely, there is no evidence that Cx. pipiens has contributed to the transmission of viruses such as DENV and CHIKV. Furthermore, experimental evidence overwhelmingly indicates that Cx. pipiens is refractory to Zika virus transmission, although some results are conflicting.

1.1.1. Emergence of West Nile Virus in Europe

West Nile virus causes a febrile illness in both humans and horses that generally resolves without complications. In a small proportion of cases (usually <1%), infected individuals develop more serious clinical symptoms and signs including encephalitis, meningitis and paralysis, followed by death in severe cases. The first isolation of WNV from the West Nile district of Uganda by Smithburn and co-workers is well documented. Serum prepared from a blood sample taken from a febrile individual was inoculated into mice from which virus was isolated. Following its discovery, research in the 1950s in Africa identified an enzootic transmission cycle involving multiple bird species as natural reservoirs for the virus and mosquitoes as the primary vector group.

The majority of countries in sub-Saharan Africa have reported evidence of WNV presence, either through isolation of the virus or through seroprevalence studies. North African countries including Egypt, Morocco and Algeria have also reported evidence of WNV. Due to its association with birds, avian migratory behaviour has been considered the most likely means for the translocation of WNV from Africa to northern latitudes. Repeated emergence of WNV in Israel, Italy, France and Spain correspond to bottlenecks of major flyways of migratory birds travelling north as they avoid the Mediterranean Sea, or cross at its narrowest points.

West Nile virus has caused sporadic outbreaks of disease in southern Europe in both humans and horses. Phylogenetic analysis has demonstrated at least eight distinct lineages of WNV, of which two, lineages 1 and 2, circulate in Europe. The most significant epidemic occurred in south eastern Romania with a focus in the capital, Bucharest. Almost 400 cases of encephalitis and 17 deaths were reported in 1996, with further cases reported in subsequent years. The Danube delta was considered the potential site of introduction but with transmission in an urban setting. Interest in WNV was also stimulated by its emergence in North America in 1999, initiating a major epidemic outbreak and highlighting its epidemic potential in other countries. The virus caused numerous cases of disease in birds, particularly North American crows (Corvus brachyrhynchos), domestic horses and humans. A wide range of indigenous mosquito species were found to be infected with WNV, however, the Cx. pipiens complex was considered to be the principal vector.

During the first decade of the 21st century, there was an increase in the number of detections of WNV outbreaks in Europe. It remains unclear if this phenomenon was due to more frequent annual re-introduction of WNV or a greater focus on surveillance in the Mediterranean Basin. The successful overwintering of virus in mosquito populations in Italy between 2008 and 2011 was a notable epidemiological shift. The virus causing the outbreaks in Italy and in other countries was identified as belonging to WNV lineage 1. West Nile virus lineage 2 was first detected in Hungary during 2004 and then spread west and south, reaching Greece prior to 2010. The outbreak in Greece was notable for involving a high number of human cases, including 33 deaths attributed to the infection. Mosquito species from the genus Culex, Aedes and other mosquito genera have been shown to transmit WNV under experimental conditions but the epidemiological significance for natural virus transmission is unclear.

1.1.2. Emergence of Usutu Virus in Europe

Usutu virus was first detected in Europe in dead blackbirds (Turdus merula) collected following a wild bird die-off event in Tuscany, Italy, in 1996. A later emergence in 2001 is better documented due to a highly visible die-off of birds around Vienna, Austria. Submission of blackbirds, great gray owls (Strix nebulosa) and a barn swallow (Hirundo rustica) resulted in detection of virus by histopathology and reverse transcription polymerase chain reaction (RT-PCR). Usutu virus has emerged in countries across southern Europe and it has subsequently spread north across western and central Europe. A small but growing number of documented cases of human infection with USUV have been recorded, although these have often been in patients with additional underlying health conditions. However, most cases of USUV infection appear to be asymptomatic.

Phylogenetic analysis using complete USUV genomes suggests that there have been multiple introductions of the virus into Europe over the past 50 years and that migrating birds are the most likely mechanism of translocation over long and short distances. Culex pipiens originating from a colony established in the Netherlands have been shown experimentally to be highly susceptible to infection with USUV when compared to WNV, although the ecoform status of the mosquitoes used was not explored.

1.1.3. Sindbis Virus in Europe

Sindbis virus (SINV) was first isolated from a pool of Cx. pipiens and/or Culex univittatus (Theobald, 1901) mosquitoes collected from the Sindbis health district, 30 km north of Cairo, Egypt. Infection causes a rash and long-lasting polyarthritis that has been recognised in northern Europe for decades. It is known colloquially as Ockelbo disease in Sweden, Pogosta disease in Finland and Karellian fever in Russia. In South Africa it has been reported to cause disease in horses; SINV infection in horses or other domestic animals has not been observed in Europe, possibly due to a lack of surveillance. The virus circulates between birds and mosquitoes with occasional spill over into human populations. Phylogenetic analysis of SINV suggests that there is long distance translocation of the virus, possibly through bird migration.

Experimental studies have shown that a range of mosquito species present in Scandinavia are capable of transmitting SINV but that Culex torrentium (Martini, 1925) demonstrated higher infection and transmission rates than Cx. pipiens. Subsequent field studies have shown higher rates of SINV infection in wild caught Cx. torrentium than in other species and this is now considered the most important vector species. Although Cx. torrentium is found across Europe and the Middle East, few cases of SINV are reported outside of northern Europe, and are limited to occasional virus isolations. The susceptibility to infection of Cx. torrentium for WNV or USUV has not been defined.

1.1.4. Other Viruses Transmitted by Culex Mosquitoes

The other Culex-transmitted viruses detected in Europe include Lednice virus (LEDV) and Rabensberg virus (RABV). LEDV, a bunyavirus, was isolated from Culex modestus (Ficalbi, 1889) in the Czech Republic in 1963; this mosquito remains the only known vector. RABV is a more recent isolation from the Czech Republic and is a virus related to WNV. It was first isolated from pools of Cx. pipiens collected in 1997 from South Moravia near the border with Austria. Batai virus (BATV) was originally detected in Culex gelidus (Theobald, 1901) in Malaysia in 1955, but in Europe it has been associated with Anopheline species.

1.2. Culex pipiens Taxonomy

The taxonomy of the Cx. pipiens complex remains a much debated subject due to the morphological similarity between some species and the varied behaviours exhibited within species. The first description of Cx. pipiens is attributed to Carl Linnaeus in 1758. The complex (or assemblage) of species includes Cx. pipiens, Cx. quinquefasciatus (Say, 1823), Cx. australicus (Dobrotworsky and Drummond, 1953) and Cx. globocoxitus (Dobrotworsky, 1953) with varied geographical distribution that has been modified by the translocation of species between continents. Additionally, some authors include the sibling species Cx. torrentium in taxonomic studies of the complex owing to its similar morphology and larval ecology. Within the species Cx. pipiens there are two ecoforms (sometimes called biotypes) recognised, pipiens (L.) and molestus (Forskål, 1775), based primarily on ecological and behavioural traits. The term molestus was first introduced by Petrus Forskål who recognised the species during an expedition to Egypt and the Arabian Peninsula. The behavioural and physiological traits reported as broadly separating the two forms are summarised in Table 2.

Evidence from several studies of European Cx. pipiens populations has indicated that ecoform molestus is a distinct species separate from ecoform pipiens and arose from a single speciation event. This contrasts with the alternative theory that molestus populations arose from repeated and independent colonisations of underground habitats by aboveground pipiens populations; other studies have shown equivocal results.

Herein, we use the following terms: (1) “Cx. pipiens complex” when referring to the group as a whole, (2) “Cx. pipiens” when referring to specimens separated from Cx. torrentium but no further, (3) “pipiens” and “molestus” in reference to the ecoforms, and (4) “pipiens/molestus” and “pipiens/quinquefasciatus” in reference to hybrid forms where appropriate.

1.3. Delineation of Species, Ecoforms and Hybrids

Differences in the structure of the male genitalia can be used to distinguish members of the complex. However, the lack of distinguishing morphological features to separate females adds complication to the identification of surveillance trap catches where females are usually the target. The presence or absence of behavioural traits such as autogeny (Table 2) have been used to identify between the forms; however, this approach is not a consistently reliable method for separating the ecoforms. Furthermore, demonstrating autogeny in wild-caught populations is labour intensive, requiring the collection and rearing of larvae, and is therefore impractical for large scale screening. This has led to the development of several molecular techniques for differentiating the two ecoforms and their hybrids (Table 3).

Initial differentiation techniques were aimed at identifying polymorphisms at 20 loci in order to differentiate above and belowground breeding populations associated with the London Underground, and to examine gene flow. This method was developed to include sequence comparison of up to 11 concatenated sequences to enable phylogenetic distinction of the two ecoforms. An alternative approach compared polymorphic microsatellite markers amplified to generate fingerprints for autogenous and anautogenous populations. Subsequent methodologies have largely been based on the polymerase chain reaction (PCR), DNA sequencing or restriction fragment length polymorphism (RFLP) (Table 3). Many of these have focused on a single locus to distinguish between the two forms, particularly the CQ11 locus (Figure 1). This end-point PCR approach is often preceded by the use of a multiplex PCR to separate Cx. torrentium from Cx. pipiens (Figure 1), although identification via comparative wing morphometrics can be used for this. In a further modification, fluorescent probes have been developed that selectively bind to the polymorphisms within the same real-time PCR amplification. Some authors have expressed caution in using only a single diagnostic marker for the identification of the Cx. pipiens complex, and advocate the use of multiple targets for maximum taxonomic clarity. For example, although a nucleotide substitution from G to A at the 3rd position of the 68th codon of the COI gene was reported as being diagnostic for form molestus over form pipiens, this finding was not replicated in a subsequent UK study that targeted the same region. In an attempt to avoid differences between assays, a recent study employed a combined four-point approach to characterising Mediterranean Cx. pipiens populations, using assays targeting the CQ11, ace-2, COI and Wolbachia (wPip) infection typing markers.

An alternative approach to species delineation is the application of matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF MS). While this application is still in relative infancy, it has been used for the identification of various vector groups and in future it may be possible to use this to define ecoforms of Cx. pipiens based on changes in protein expression.

1.4. Distribution and Hybridisation

Culex pipiens is widely distributed across Eurasia and further afield. Our understanding of the local and regional distribution of its ecoforms has, however, developed only relatively recently, aided by the increasing use of molecular species delineation methods. There remains, however, a poor understanding of the relationship between the genetics of the ecoforms and their phenotype. Initial evidence indicated a fairly consistent separation between the habitats of each ecoform: the ubiquitous pipiens ecoform was associated with natural and artificial aboveground habitats across rural and urban areas and the molestus form was found in urban underground habitats. Particularly in northern Europe, this habitat distinction was believed to serve as a barrier to hybridisation between the forms and this was supported by limited success in breeding between forms under laboratory conditions.

Present evidence, however, suggests that this habitat separation is far less rigid, with cross-breeding experiments and analysis of genetic markers from field and colony specimens indicating that inter-breeding populations of pipiens and molestus can be found sympatrically in both above- and belowground urban habitats, as well as in rural and semi-rural areas. Indeed, natural hybrid pipiens/molestus forms have now been reported from at least 12 European countries (Figure 2a) with reported rates of hybridisation of up to 25.7%. The relative abundance of each of the forms and hybridisation rates have been found to vary across latitudes, with the proportion of molestus populations relative to pipiens increasing from northern to southern latitudes. To add further complexity, hybridisation of Cx. pipiens with Cx. quinquefasciatus has been reported from the Mediterranean Basin (Figure 2b), despite sympatric populations of these species existing without hybridisation in East Africa.

The occurrence of natural hybrid populations has important consequences for the risks of pathogen transmission. Changes to mosquito host preference, vector competence, the occurrence of autogeny and the ability to forgo diapause and continue reproduction through the winter months may all alter virus transmission dynamics. This may have contributed to the persistence of WNV in Romania during the 1990s where the presence of mosquitoes indoors and in flooded basements were considered risk factors for human infection. Additionally, the strains of the endosymbiont Wolbachia pipientis associated with Cx. quinquefasciatus and the different ecoforms of Cx. pipiens differ, and the impact of such differences on vector competence is not fully understood. Studies of Cx. pipiens populations in Portugal demonstrated that gene flow occurred predominantly from the molestus to the pipiens form. Asymmetric gene flow in this fashion could alter feeding preferences of Cx. pipiens from an ornithophilic to mammalophilic feeding preference, as demonstrated in the USA. The vector competence of molestus populations to WNV in The Netherlands was lower (6–10%) than that of pipiens (0–32%) and hybrid (0–14%) forms. In this context, gene flow from pipiens to molestus could result in increased vector competence and thus may be equally important in influencing local pathogen transmission dynamics.

Culex torrentium has also been reported from many countries across Europe (Figure 3) where its larvae are often found in sympatry with Cx. pipiens. In many studies little morphological separation is performed, thus masking the true distribution of the two species. Initially believed to be a rare European species, Cx. torrentium is now recognised to be widespread in northern and central regions of Europe. When compared with Cx. pipiens, these species form an apparent contrasting gradient of abundance: in northern regions Cx. torrentium dominates, in central Europe both species exist in similar proportions, and in southern Europe Cx. pipiens is the dominant species and Cx. torrentium is rarely reported. The current distribution of Cx. torrentium may reflect a range expansion, perhaps in response to favourable anthroponotic environmental changes, but the misidentification of females as Cx. pipiens prior to the widespread use of molecular analyses may have hindered information on its distribution.

1.5. Culex pipiens Blood-Feeding Behaviour

A critical behavioural trait relevant to arthropod-borne virus transmission is a vector’s host feeding pattern. Host selection determines the exposure of a mosquito to pathogens and its involvement in enzootic, zoonotic or anthroponotic transmission cycles. Host selection by mosquitoes is a complex phenomenon, influenced by an interplay of genetic and environmental factors. The latter includes the local and seasonal presence of vertebrate hosts, host defensive behaviour against biting and the presence of pathogens in the arthropod, host, or both, which may influence rates of vector-host contact. Evidence for preferential feeding on specific hosts may be derived from studies that identify the blood meal hosts of wild-caught engorged mosquitoes, or semi-field or laboratory tests offering a choice of feeding from different hosts.

The pipiens ecoform is considered to be almost exclusively ornithophilic (bird-feeding), whilst the molestus ecoform feeds on other mammalian hosts, including humans (Table 2). Here, we collated data from 29 European studies identifying the blood meals of Cx. pipiens (Table 4).

Collectively, these data show feeding of Cx. pipiens on a wide range of hosts encompassing mammals (eight orders, 12 families and 17 species), birds (14 orders, 33 families, 82 species) and reptiles (two orders, three families, three species). Eight of these studies identified specimens to ecoform, and three of these successfully collected blood-fed specimens of both ecoforms and their hybrids, identified by sequence analysis of the CQ11 locus. Collectively, these latter three studies identified both ecoforms and their hybrids as feeding on both mammals and birds. Interestingly, all found that birds were highly utilised by the pipiens and molestus ecoforms plus their hybrids (Figure 4), with no significant differences in feeding preference between the forms. These results contrast with findings in the USA showing that specimens with a higher proportion of molestus ancestry fed more frequently on humans. Reasons for these disparate findings may lie with geographic or seasonal differences in host availability, the relatively low sample sizes inherent with the challenges of collecting blood-fed specimens, or with differences in the microsatellite markers used to identify the forms in each study.

Relatively few manipulative comparisons of host selection, whereby mosquitoes are offered choices to feed on different hosts, have been carried out with Cx. pipiens under field, semi-field or laboratory conditions. Preferential attraction was recorded towards chicks by the pipiens ecoform, to humans by the molestus ecoform, and intermediate feeding behaviour in pipiens/molestus hybrids from field-collected populations in Chicago, USA. Choice tests can be an effective method to compare feeding preferences between individual hosts, but to our knowledge, these have not been conducted to compare the ecoforms and hybrids of European populations of Cx. pipiens.

Field studies collecting mosquitoes attempting to feed on live human or animal baits can also greatly contribute to our understanding of host preference. Several field studies have reported human-biting Cx. pipiens; studies in Portugal and the UK collected both pipiens and molestus ecoforms by human landing catch. Although the study in Portugal identified human blood in one engorged pipiens female, the specimens collected by human landing catch in both studies did not contain blood to permit confirmation of human feeding. However, this collection method is considered the gold-standard approach for assessing mosquito-human contact rates, with mosquito feeding (or at least probing) assumed to occur after landing. Combining these field data with laboratory choice tests and, although challenging, with blood meal studies that are coupled with comprehensive surveys of vertebrate hosts in the sample area to assess the impact of host availability, will contribute further to our understanding of host selection and preference of members of the Culex pipiens complex. However, studies where wild mosquitoes are offered a choice of host are very rare and findings such as those reported above could therefore represent opportunistic feeding rather than a true preference.

2. Future Research Directions

Our understanding of the Cx. pipiens complex has expanded rapidly in recent years, but there remain many intriguing and as yet unexplored questions concerning their biology and ecology. Below we highlight four areas of research important to defining the impact of Cx. pipiens on present and future virus transmission in Europe.

(1) What factors lead to successful arbovirus transmission by populations of Culex pipiens?

The distribution of Culex-transmitted arboviruses is not uniform across Europe. Identification of the different factors that lead to successful transmission of viruses and those that preclude virus emergence are critical to understanding this distribution. Northern Europe has seasonally abundant populations of Cx. pipiens that appear to support transmission of USUV but not WNV. This suggests environmental and climatic factors alone cannot explain the absence of WNV from countries such as Germany, Poland, The Netherlands and the United Kingdom. In North America, Culex species, including Cx. pipiens, enabled rapid spread of West Nile virus across the continent with no apparent barriers. Expanding upon recent work investigating the distribution of the ecoforms of Cx. pipiens across Europe is essential to furthering our understanding of the relationship between the ecoforms and their hybrids with current arbovirus distribution patterns. Furthermore, as many important arboviruses exist in bird-mosquito-bird transmission cycles, identifying hotspots of high mosquito and resident and migratory bird populations will enable better targeting of interventions in advance of a novel virus introduction. Such hotspots may include rural wetland areas but could, increasingly, include more urbanised areas. At the level of the mosquito, there remain many questions regarding the complex interplay of genetic and environmental factors that influence vector competence and mosquito-virus-host interactions. These include the extrinsic incubation period, viral adaptivity, mosquito and host immunity and mosquito behaviour. In reference to the latter, newly-emerged Australian ecoform molestus females preferentially delay blood-feeding until after laying their first egg batch. If such high levels of obligatory autogeny exist in European populations, this would not only provide a highly beneficial population survival mechanism but may influence the transovarial maintenance of virus through several generations. Finally, the survival of virus in overwintering Cx. pipiens is likely a critical factor involved in the maintenance of transmission cycles in Europe; a recent study detected WNV RNA in overwintering Cx. pipiens in the Czech Republic. Further investigation of the factors influencing overwintering survival, post-hibernation emergence, and subsequent dispersal of Cx. pipiens and its ecoforms, as conducted elsewhere, will improve our understanding of the role of overwintering in virus maintenance, particularly in regions of Europe that experience colder winters.

(2) What are the potential impacts of a changing environment?

That climate changes are occurring and will impact both native and non-native arthropod fauna worldwide is well established. The potential influences on arthropod-borne pathogens have been explored, although the specific effects will vary considerably according to mosquito species biology and the region concerned. Anthroponotic changes influencing the structure of the environment may be equally important in altering mosquito populations at the local or regional scale. For example, the creation of urban wetlands as part of sewage treatment works could increase available eutrophic habitat particularly suitable for ecoform molestus. Increasing urbanisation could provide additional container habitats suitable for existing urban mosquito populations, or facilitate an adaptive shift by other species towards the utilisation of urban habitats, as evidenced by an increasing urban population of Anopheles plumbeus (Stephens, 1828) in various parts of north-western Europe. Urban centres could be at further risk of vector-borne disease if existing temperature rises were compounded by the urban heat island effect in such locations, although the precise effects of this phenomenon on pathogen transmission risk are likely to be complex. The storage of water during periods of drought could additionally provide increased urban habitat for mosquito breeding, whilst the reversion of arable land to wetlands could provide further habitat for Culex mosquitoes and provide a location where grazing animals come into contact with migratory birds.

(3) What are the key factors influencing rates of hybridisation?

The variable rates of hybridisation in European populations between sympatric populations of pipiens and molestus ecoforms indicate the existence of multiple barriers to hybridisation that extend beyond simple allopatric reproductive isolation. Although in parts of Europe hybridisation rates are low, rates in southern Europe may approach those reported from northern Africa. To what extent reproductive barriers are behavioural, such as environmental requirements for swarm formation or specificity of matched wing beat frequencies, or intrinsic, for example mediated by commensal Wolbachia strains and cytoplasmic incompatibility, is currently unknown. Furthermore, although human-mediated transport of mosquitoes may facilitate long-distance species translocation and provide opportunities for hybridisation aboveground, to what extent are belowground molestus populations able to disperse within and beyond their existing habitats? Approaches such as the use of mark-release-recapture aided by fluorescently- or immune-marked insects in belowground systems could, for example, reveal the dispersal potential of form molestus.

(4) How do the olfactory responses to semiochemicals of host and environmental origin differ?

Furthering the understanding of the responses of the Cx. pipiens complex to volatile compounds produced by vertebrate hosts, nectar sources and larval habitats will facilitate the development of novel repellents, attractants and more optimal approaches to surveillance and control. To date, the olfactory responses of Culex species to host odours have been investigated for Cx. quinquefasciatus, and to flower odours in ecoform pipiens and molestus. However, directly comparative studies of the olfactory responses between the ecoforms have not been conducted, and paired trap comparison studies comparing above- and belowground collections remain unexplored. Recent work has shown that ecoforms pipiens and molestus, plus their hybrids, were collected in similar ratios by BG-Sentinel and Mosquito Magnet Liberty Plus traps. However, Cx. torrentium was found to be under-represented in CDC light trap catches in Germany and Sweden in comparison to Cx. pipiens and although the authors did not molecularly identify specimens to ecoform, these results illustrate the need for further field investigation using other trap types.

In summary, it is vital that data on members of the Cx. pipiens complex is collected from countries across Europe and at a range of geographic scales that reflect different ecological zones. Comparisons should also be made between urban and rural populations and those in intermediate areas. Habitat differences may be more important in influencing distribution and hybridisation rates than broader latitudinal trends. Studies conducted at the regional, national and pan-European level will provide critical data to model trends in mosquito biology and virus transmission, and to better inform regional approaches to surveillance and control. However, these large-scale studies cannot replace targeted field-based studies which are critical to understand the factors influencing transmission at the level of the vector and its hosts in different local habitats. Finally, although these research questions span several fields, it has become increasingly clear that future studies should, insofar as is possible, identify Cx. pipiens to the level of both species and ecoform. The continued decrease in costs and increase in the speed of molecular identification approaches will no doubt greatly contribute towards this goal.

3. Conclusions

Current evidence from across Europe highlights the importance of the Cx. pipiens complex in the current and potential future transmission of important medical and veterinary arboviruses. It is therefore imperative that a concerted effort be made between research and governmental agencies across Europe to better target future sampling efforts to answer the remaining questions concerning the ecology and genetics of mosquito and pathogen that influence this association.

Surveillance for mosquito-borne viruses in mosquito populations varies widely across Europe. Extensive surveillance is conducted in northern Italy where cases of WNV occur annually in an attempt to detect virus in mosquitoes populations. This offers the opportunity for public health authorities to warn health professionals before the occurrence of human disease. Both Germany and Switzerland conduct extensive surveillance to detect invasive mosquitoes and the emergence of virus infections. This has proven useful in mapping the spread of USUV across Europe. In the majority of countries across Europe, however, surveillance is reactive in response to disease outbreaks or changes in the mosquito population.

The extent of the distribution of the specific forms of Cx. pipiens is just beginning to be defined. However, evidence indicates that latitudinal differences in the distribution of Cx. pipiens forms and their hybrids, together with the distribution of the sibling species Cx. torrentium, may influence the transmission dynamics of arboviruses in Europe. However, the picture is more complicated than simply this fact and will include the effect of different environmental conditions on the life cycle and behaviour of the mosquitoes, as well as intrinsic factors such as vector competence. In addition, despite the importance of this species in current and potential pathogen transmission, increasing our understanding of how species complexes as a whole function within an ecosystem to contribute to pathogen transmission is vitally important. For example, Rift Valley fever virus outbreaks involve multiple species that act sequentially depending on environmental circumstances. Therefore, maintenance of surveillance approaches that target a wide range of mosquito species should be used.

Current evidence continues to support the importance of birds as a major blood-meal host for Cx. pipiens across Europe. However, there is considerable evidence from blood meal and host-baited studies that ecoform pipiens can also take blood meals from humans and other mammals. Conversely, ecoform molestus also feeds to a considerable extent on birds, in many cases to the same degree as the pipiens ecoform. Therefore, it may be necessary to take a broader view and consider the potential for both ecoforms to act as enzootic and bridge vectors of medically important arboviruses.

References 224

  1. 1.

    Gratz N.G. The Vector- and Rodent-Borne Diseases of Europe and North America: Their Distribution and Public Health Burden. 2006.

    Not in our library

  2. 2.

    Harbach R.E. Mosquito Taxonomic Inventory.

    Not in our library

  3. 3.

    Harbach R.E., Howard T.M. Index of currently recognized mosquito species (Diptera: Culicidae). Eur. Mosq. Bull. 2007;23:1-66.

    Not in our library

  4. 4.

    Medlock J.M., Hansford K.M., Versteirt V., Cull B., Kampen H., Fontenille D., Hendrickx G., Zeller H., van Bortel W., Schaffner F. An entomological review of invasive mosquitoes in Europe. Bull. Entomol. Res. 2015;1-27.

    Not in our library10.1017/s0007485315000103

  5. 5.

    Adhami J., Reiter P. Introduction and establishment of Aedes (Stegomyia) albopictus Skuse (Diptera: Culicidae) in Albania. J. Am. Mosq. Control Assoc. 1998;14:340-343.

    Not in our library

  6. 6.

    Sabatini A., Raineri V., Trovato G., Coluzzi M. Aedes albopictus in Italy and possible diffusion of the species into the Mediterranean area. Parassitologia. 1990;32:301-304.

    Not in our library

  7. 7.

    European Centre for Disease Prevention and Control. Aedes Albopictus—Factsheet for Experts. 2016.

    Not in our library

  8. 8.

    Margarita Y., Grácio A.J., Lencastre I., Silva A.C., Novo M.T., Sousa C.A. First record of Aedes (Stegomyia) aegypti (Linnaeus, 1762) (Diptera, Culicidae) in Madeira Island-Portugal. Acta Parasitológica Port. 2006;13:59-61.

    Not in our library

  9. 9.

    Akiner M.M., 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:e0004664.

    Not in our library10.1371/journal.pntd.0004664

  10. 10.

    Versteirt V., de Clercq E.M., Fonseca D.M., Pecor J., Schaffner F., Coosemans M., van Bortel W. Bionomics of the established exotic mosquito species Aedes koreicus in Belgium, Europe. J. Med. Entomol. 2012;49:1226-1232.

    Not in our library10.1603/me11170

  11. 11.

    Capelli G., Drago A., Martini S., Montarsi F., Soppelsa M., Delai N., Ravagnan S., Mazzon L., Schaffner F., Mathis A., et al. First report in Italy of the exotic mosquito species Aedes (Finlaya) koreicus, a potential vector of arboviruses and filariae. Parasit. Vectors. 2011;4:188.

    Not in our library10.1186/1756-3305-4-188

  12. 12.
  13. 13.

    Calzolari M., Bonilauri P., Bellini R., Albieri A., Defilippo F., Maioli G., Galletti G., Gelati A., Barbieri I., Tamba M., et al. Evidence of simultaneous circulation of West Nile and Usutu viruses in mosquitoes sampled in Emilia-Romagna region (Italy) in 2009. PLoS ONE. 2010;5:e14324.

    Not in our library10.1371/journal.pone.0014324

  14. 14.

    Papa A., Xanthopoulou K., Gewehr S., Mourelatos S. Detection of West Nile virus lineage 2 in mosquitoes during a human outbreak in Greece. Clin. Microbiol. Infect. 2011;17:1176-1180.

    Not in our library10.1111/j.1469-0691.2010.03438.x

  15. 15.

    Savini G., Capelli G., Monaco F., Polci A., Russo F., Di Gennaro A., Marini V., Teodori L., Montarsi F., Pinoni C., et al. Evidence of West Nile virus lineage 2 circulation in Northern Italy. Vet. Microbiol. 2012;158:267-273.

    Not in our library10.1016/j.vetmic.2012.02.018

  16. 16.

    Almeida A.P.G., Freitas F.B., Novo M.T., Sousa C.A., Rodrigues J.C., Alves R., Esteves A. Mosquito surveys and West Nile virus screening in two different areas of southern Portugal, 2004–2007. Vector-Borne Zoonotic Dis. 2010;10:673-680.

    Not in our library10.1089/vbz.2009.0245

  17. 17.
  18. 18.

    Fortuna C., Remoli M.E., Di Luca M., Severini F., Toma L., Benedetti E., Bucci P., Montarsi F., Minelli G., Boccolini D., et al. Experimental studies on comparison of the vector competence of four Italian Culex pipiens populations for West Nile virus. Parasit. Vectors. 2015;8:463.

    Not in our library10.1186/s13071-015-1067-z

  19. 19.

    Fros J.J., Miesen P., Vogels C.B., Gaibani P., Sambri V., Martina B.E., Koenraadt C.J., van Rij R.P., Vlak J.M., Takken W., et al. Comparative Usutu and West Nile virus transmission potential by local Culex pipiens mosquitoes in north-western Europe. One Heal. 2015;1:31-36.

    Not in our library10.1016/j.onehlt.2015.08.002

  20. 20.
  21. 21.

    Farajollahi A., Fonseca D.M., Kramer L.D., Kilpatrick A.M. “Bird biting” mosquitoes and human disease: A review of the role of Culex pipiens complex mosquitoes in epidemiology. Infect. Genet. Evol. 2011;11:1577-1585.

    Not in our library10.1016/j.meegid.2011.08.013

  22. 22.

    Vinogradova E.B. Culex pipiens pipiens mosquitoes: Taxonomy, distribution, ecology, physiology, genetics, applied importance and control. Pensoft Ser. Parasitol. 2000;2:250.

    Not in our library

  23. 23.

    Lundström J.O. Mosquito-borne viruses in western Europe: A review. J. Vector Ecol. 1999;24:1-39.

    Not in our library

  24. 24.

    Hubálek Z. Mosquito-borne viruses in Europe. Parasitol. Res. 2008;103:S29-S43.

    Not in our library10.1007/s00436-008-1064-7

  25. 25.

    Snow K.R., Medlock J.M. The potential impact of climate change on the distribution and prevalence of mosquitoes in Britain. Eur. Mosq. Bull. 2006;21:1-10.

    Not in our library

  26. 26.

    Becker N. Influence of climate change on mosquito development and mosquito-borne diseases in Europe. Parasitol. Res. 2008;103:19-28.

    Not in our library10.1007/s00436-008-1210-2

  27. 27.

    Gould E.A., Higgs S. Impact of climate change and other factors on emerging arbovirus diseases. Trans. R. Soc. Trop. Med. Hyg. 2009;103:109-121.

    Not in our library10.1016/j.trstmh.2008.07.025

  28. 28.

    Aliota M.T., Peinado S.A., Osorio J.E., Bartholomay L.C. Culex pipiens and Aedes triseriatus mosquito susceptibility to Zika Virus. Emerg. Infect. Dis. 2016;22:1857-1859.

    Not in our library10.3201/eid2210.161082

  29. 29.

    Kenney J.L., Romo H., Duggal N.K., Tzeng W.P., Burkhalter K.L., Brault A.C., Savage H.M. Transmission incompetence of Culex quinquefasciatus and Culex pipiens pipiens from North America for Zika virus. Am. J. Trop. Med. Hyg. 2017;96:1235-1240.

    Not in our library10.4269/ajtmh.16-0865

  30. 30.

    Heitmann A., Jansen S., Lühken R., Leggewie M., Badusche M., Pluskota B., Becker N., Vapalahti O., Schmidt-Chanasit J., Tannich E. Experimental transmission of zika virus by mosquitoes from central Europe. Eurosurveillance. 2017;22:30437.

    Not in our library10.2807/1560-7917.es.2017.22.2.30437

  31. 31.

    Amraoui F., Atyame-Nten C., Vega-Rúa A., Lourenço-De-Oliveira R., Vazeille M., Failloux A.B. Culex mosquitoes are experimentally unable to transmit zika virus. Eurosurveillance. 2016;21:30333.

    Not in our library10.2807/1560-7917.es.2016.21.35.30333

  32. 32.

    Boccolini D., Toma L., Di Luca M., Severini F., Romi R., Remoli M.E., Sabbatucci M., Venturi G., Rezza G., Fortuna C. Experimental investigation of the susceptibility of Italian Culex pipiens mosquitoes to zika virus infection. Eurosurveillance. 2016;21:30328.

    Not in our library10.2807/1560-7917.es.2016.21.35.30328

  33. 33.
  34. 34.
  35. 35.
  36. 36.

    Huang Y.-J.S., Ayers V.B., Lyons A.C., Unlu I., Alto B.W., Cohnstaedt L.W., Higgs S., Vanlandingham D.L. Culex species mosquitoes and Zika virus. Vector-Borne Zoonotic Dis. 2016;16:673-676.

    Not in our library10.1089/vbz.2016.2058

  37. 37.
  38. 38.
  39. 39.

    Medlock J.M., Snow K.R., Leach S.A. Possible ecology and epidemiology of medically important mosquito-borne arboviruses in Great Britain. Epidemiol. Infect. 2007;135:466-482.

    Not in our library10.1017/s0950268806007047

  40. 40.
  41. 41.

    Tomasello D., Schlagenhauf P. Chikungunya and dengue autochthonous cases in Europe, 2007–2012. Travel Med. Infect. Dis. 2013;11:274-284.

    Not in our library10.1016/j.tmaid.2013.07.006

  42. 42.

    Pecorari M., Longo G., Gennari W., Grottola A., Sabbatini A., Tagliazucchi S., Savini G., Monaco F., Simone M.L., Lelli R., et al. First human case of Usutu virus neuroinvasive infection, Italy, August–September 2009. Euro. Surveill. 2009;14:19446.

    Not in our library

  43. 43.
  44. 44.

    Campbell G.L., Marfin A.A., Lanciotti R.S., Gubler D.J. West Nile virus. Lancet Infect. Dis. 2002;2:519-529.

    Not in our library10.1016/s1473-3099(02)00368-7

  45. 45.
  46. 46.

    Smithburn K.C., Hughes T.P., Burke A.W., Paul J.H. A neurotropic virus isolated from the blood of a native of Uganda. Am. J. Trop. Med. Hyg. 1940;20:471-492.

    Not in our library10.4269/ajtmh.1940.s1-20.471

  47. 47.

    Work T.H., Hurlbut H.S., Taylor R.M. Indigenous wild birds of the Nile-delta as potential West Nile virus circulating reservoirs. Am. J. Trop. Med. Hyg. 1955;4:872-888.

    Not in our library10.4269/ajtmh.1955.4.872

  48. 48.

    Cabre O., Grandadam M., Marié J.-L., Gravier P., Prangé A., Santinelli Y., Rous V., Bourry O., Durand J.-P., Tolou H., et al. West Nile Virus in horses, sub-Saharan Africa. Emerg. Infect. Dis. 2006;12:1958-1960.

    Not in our library10.3201/eid1212.060042

  49. 49.

    Melnick J.L., Paul J.R., Riordan J.T., Barnett V.H., Goldblum N., Zabin E. Isolation from human sera in Egypt of a virus apparently identical to West Nile Virus. Proc. Soc. Exp. Biol. Med. 1951;77:661-665.

    Not in our library10.3181/00379727-77-18884

  50. 50.
  51. 51.

    Lafri I., Prat C.M., Bitam I., Gravier P., Besbaci M., Zeroual F., Ben-Mahdi M.H., Davoust B., Leparc-Goffart I. Seroprevalence of West Nile virus antibodies in equids in the North-East of Algeria and detection of virus circulation in 2014. Comp. Immunol. Microbiol. Infect. Dis. 2017;50:8-12.

    Not in our library10.1016/j.cimid.2016.11.005

  52. 52.

    Rappole J.H., Hubálek Z. Migratory birds and West Nile virus. J. Appl. Microbiol. 2003;94:47S-58S.

    Not in our library10.1046/j.1365-2672.94.s1.6.x

  53. 53.
  54. 54.

    Nikolay B. A review of West Nile and Usutu virus co-circulation in Europe: How much do transmission cycles overlap? Trans. R. Soc. Trop. Med. Hyg. 2015;109:609-618.

    Not in our library10.1093/trstmh/trv066

  55. 55.

    Fall G., Di Paola N., Faye M., Dia M., de Melo Freire C.C., Loucoubar C., de Andrade Zanotto P.M., Faye O., Sall A.A. Biological and phylogenetic characteristics of West African lineages of West Nile virus. PLoS Negl. Trop. Dis. 2017;11:e0006078.

    Not in our library10.1371/journal.pntd.0006078

  56. 56.

    Cernescu C., Nedelcu N.I., Tardei G., Ruta S., Tsai T.F. Continued transmission of West Nile virus to humans in southeastern Romania, 1997–1998. J. Infect. Dis. 2000;181:710-712.

    Not in our library10.1086/315225

  57. 57.

    Lanciotti R.S. Origin of the West Nile virus responsible for an outbreak of encephalitis in the Northeastern United States. Science. 1999;286:2333-2337.

    Not in our library10.1126/science.286.5448.2333

  58. 58.
  59. 59.

    Hamer G., Kitron U.D., Goldberg T.L., Brawn J.D., Loss S.R., Ruiz M.O., Hayes D.B., Walker E.D. Host selection by Culex pipiens mosquitoes and West Nile virus amplification. Am. J. Trop. Med. Hyg. 2009;80:268-278.

    Not in our library

  60. 60.

    Andreadis T.G. The contribution of Culex pipiens complex mosquitoes to transmission and persistence of West Nile virus in North America. J. Am. Mosq. Control Assoc. 2012;28:137-151.

    Not in our library10.2987/8756-971x-28.4s.137

  61. 61.

    Calzolari M., Bonilauri P., Bellini R., Albieri A., Defilippo F., Tamba M., Tassinari M., Gelati A., Cordioli P., Angelini P., et al. Usutu virus persistence and West Nile virus inactivity in the Emilia-Romagna region (Italy) in 2011. PLoS ONE. 2013;8:e63978.

    Not in our library10.1371/journal.pone.0063978

  62. 62.

    Sotelo E., Fernández-Pinero J., Llorente F., Vázquez A., Moreno A., Agüero M., Cordioli P., Tenorio A., Jimeénez-Clavero M.Á. Phylogenetic relationships of Western Mediterranean West Nile virus strains (1996–2010) using full-length genome sequences: Single or multiple introductions? J. Gen. Virol. 2011;92:2512-2522.

    Not in our library10.1099/vir.0.033829-0

  63. 63.
  64. 64.

    Mani P., Rossi G., Perrucci S., Bertini S. Mortality of Turdus merula in Tuscany. Sel. Vet. 1998;8:749-753.

    Not in our library

  65. 65.
  66. 66.

    Bakonyi T., Erdélyi K., Brunthaler R., Dán Á., Weissenböck H., Nowotny N. Usutu virus, Austria and Hungary, 2010–2016. Emerg. Microbes Infect. 2017;6:e85.

    Not in our library10.1038/emi.2017.72

  67. 67.

    Cavrini F., Gaibani P., Longo G., Pierro A.M., Rossini G., Bonilauri P., Gerundi G.E., Di Benedetto F., Pasetto A., Girardis M., et al. Usutu virus infection in a patient who underwent orthotropic liver transplantation, Italy, August–September 2009. Eurosurveillance. 2009;14:19448.

    Not in our library

  68. 68.

    Santini M., Vilibic-Cavlek T., Barsic B., Barbic L., Savic V., Stevanovic V., Listes E., Di Gennaro A., Savini G. First cases of human Usutu virus neuroinvasive infection in Croatia, August–September 2013: Clinical and laboratory features. J. Neurovirol. 2014;21:92-97.

    Not in our library10.1007/s13365-014-0300-4

  69. 69.

    Cadar D., Maier P., Müller S., Kress J., Chudy M., Bialonski A., Schlaphof A., Jansen S., Jöst H., Tannich E., et al. Blood donor screening for west nile virus (WNV) revealed acute usutu virus (USUV) infection, Germany, September 2016. Eurosurveillance. 2017;22:30501.

    Not in our library10.2807/1560-7917.es.2017.22.14.30501

  70. 70.

    Taylor R.M., Hurlbut H.S., Work T.H., Kingston J.R., Frothingham T.E. Sindbis virus: A newly recognized arthropod-transmitted virus. Am. J. Trop. Med. Hyg. 1955;4:844-862.

    Not in our library10.4269/ajtmh.1955.4.844

  71. 71.

    Espmark A., Niklasson B. Ockelbo disease in Sweden: epidemiological, clinical, and virological data from the 1982 outbreak. Am. J. Trop. Med. Hyg. 1984;33:1203-1211.

    Not in our library10.4269/ajtmh.1984.33.1203

  72. 72.

    Van Niekerk S., Human S., Williams J., van Wilpe E., Pretorius M., Swanepoel R., Venter M. Sindbis and Middelburg old world alphaviruses associated with neurologic disease in horses, South Africa. Emerg. Infect. Dis. 2015;21:2225-2229.

    Not in our library10.3201/eid2112.150132

  73. 73.

    Adouchief S., Smura T., Sane J., Vapalahti O., Kurkela S. Sindbis virus as a human pathogen—Epidemiology, clinical picture and pathogenesis. Rev. Med. Virol. 2016;26:221-241.

    Not in our library10.1002/rmv.1876

  74. 74.

    Lundström J.O., Pfeffer M. Phylogeographic structure and evolutionary history of Sindbis virus. Vector-Borne Zoonotic Dis. 2010;10:889-907.

    Not in our library10.1089/vbz.2009.0069

  75. 75.

    Lundström J.O., Niklasson B., Francy D.B. Swedish Culex torrentium and Cx. pipiens (Diptera: Culicidae) as experimental vectors of Okelbo virus. J. Med. Entomol. 1990;27:561-563.

    Not in our library10.1093/jmedent/27.4.561

  76. 76.

    Lundström J.O., Turell M.J., Niklasson B. Effect of environmental temperature on the vector competence of Culex pipiens and Cx. torrentium for Okelbo virus. Am. J. Trop. Med. Hyg. 1990;43:534-542.

    Not in our library10.4269/ajtmh.1990.43.534

  77. 77.
  78. 78.

    Eiden M., Ziegler U., Keller M., Müller K., Granzow H., Jöst H., Schmidt-Chanasit J., Groschup M.H. Isolation of sindbis virus from a hooded crow in Germany. Vector-Borne Zoonotic Dis. 2014;14:220-222.

    Not in our library10.1089/vbz.2013.1354

  79. 79.

    Hesson J.C., Ostman O., Schäfer M., Lundström J.O. Geographic distribution and relative abundance of the sibling vector species Culex torrentium and Culex pipiens in Sweden. Vector-Borne Zoonotic Dis. 2011;11:1383-1389.

    Not in our library10.1089/vbz.2011.0630

  80. 80.

    Málková D., Danielová V., Minár B., Rosický B., Casals J. Isolation of Yaba 1 arbovirus in Czechoslovakia. Acta Virol. 1972;16:93.

    Not in our library

  81. 81.

    Danielová V. To the problem of the vector of Lednice virus. Folia Parasitol. (Praha). 1984;31:379-382.

    Not in our library

  82. 82.
  83. 83.

    Jöst H., Bialonski A., Schmetz C., Günther S., Becker N., Schmidt-Chanasit J. Isolation and phylogenetic analysis of Batai virus, Germany. Am. J. Trop. Med. Hyg. 2011;84:241-243.

    Not in our library10.4269/ajtmh.2011.10-0483

  84. 84.

    Weitzel T., Collado A., Jöst A., Pietsch K., Storch V., Becker N. Genetic differentiation of populations within the Culex pipiens complex and phylogeny of related species. J. Am. Mosq. Control Assoc. 2009;25:6-17.

    Not in our library10.2987/08-5699.1

  85. 85.

    Becker N., Jöst A., Weitzel T. The Culex pipiens complex in Europe. J. Am. Mosq. Control Assoc. 2012;28:53-67.

    Not in our library10.2987/8756-971x-28.4s.53

  86. 86.

    Harbach R.E. Culex pipiens: Species versus species complex—taxonomic history and perspective. J. Am. Mosq. Control Assoc. 2012;28:10-23.

    Not in our library10.2987/8756-971x-28.4.10

  87. 87.

    Russell R.C. A review of the status and significance of the species within the Culex pipiens group in Australia. J. Am. Mosq. Control Assoc. 2012;28:24-27.

    Not in our library10.2987/8756-971x-28.4s.24

  88. 88.

    Fonseca D.M., Lapointe D.A., Fleischer R.C. Bottlenecks and multiple introductions: Population genetics of the vector of avian malaria in Hawaii. Mol. Ecol. 2000;9:1803-1814.

    Not in our library10.1046/j.1365-294x.2000.01070.x

  89. 89.

    Hesson J.C., Rettich F., Merdić E., Vignjević G., Ostman O., Schäfer M., Schaffner F., Foussadier R., Besnard G., Medlock J.M., et al. The arbovirus vector Culex torrentium is more prevalent than Culex pipiens in northern and central Europe. Med. Vet. Entomol. 2014;28:179-186.

    Not in our library10.1111/mve.12024

  90. 90.

    Fonseca D.M., Keyghobadi N., Malcolm C.A., Mehmet C., Schaffner F., Mogi M., Fleischer R.C., Wilkerson R.C. Emerging vectors in the Culex pipiens complex. Science. 2004;303:1535-1538.

    Not in our library10.1126/science.1094247

  91. 91.

    Shaikevich E.V. PCR-RFLP of the COI gene reliably differentiates Cx. pipiens, Cx. pipiens f. molestus and Cx. torrentium of the Pipiens Complex. Eur. Mosq. Bull. 2007;23:25-30.

    Not in our library

  92. 92.

    Byrne K., Nichols R.A. Culex pipiens in London Underground tunnels: Differentiation between surface and subterranean populations. Heredity (Edinb.). 1999;82:7-15.

    Not in our library10.1038/sj.hdy.6884120

  93. 93.

    Kothera L., Godsey M., Mutebi J.-P., Savage H.M. A comparison of aboveground and belowground populations of Culex pipiens (Diptera: Culicidae) mosquitoes in Chicago, Illinois, and New York City, New York, using microsatellites. J. Med. Entomol. 2010;47:805-813.

    Not in our library10.1603/me10031

  94. 94.

    Dobrotworsky N.V. The problem of the Culex pipiens complex in the South Pacific (including Australia). Bull. World Health Organ. 1967;37:251-255.

    Not in our library

  95. 95.

    Kent R.J., Harrington L.C., Norris D.E. Genetic differences between Culex pipiens f. molestus and Culex pipiens pipiens (Diptera: Culicidae) in New York. J. Med. Entomol. 2007;44:50-59.

    Not in our library10.1093/jmedent/41.5.50

  96. 96.

    Keyghobadi N., Matrone M.A., Ebel G.D., Kramer L.D., Fonseca D.M. Microsatellite loci from the northern house mosquito (Culex pipiens), a principal vector of West Nile virus in North America. Mol. Ecol. Notes. 2004;4:20-22.

    Not in our library10.1046/j.1471-8286.2003.00557.x

  97. 97.

    Bahnck C.M., Fonseca D.M. Rapid assay to identify the two genetic forms of Culex (Culex) pipiens L. (Diptera: Culicidae) and hybrid populations. Am. J. Trop. Med. Hyg. 2006;75:251-255.

    Not in our library

  98. 98.

    Smith J.L., Fonseca D.M. Rapid assays for identification of members of the Culex (Culex) pipiens complex, their hybrids, and other sibling species (Diptera: Culicidae). Am. J. Trop. Med. Hyg. 2004;70:339-345.

    Not in our library

  99. 99.

    Börstler J., Lühken R., Rudolf M., Steinke S., Melaun C., Becker S., Garms R., Krüger A. The use of morphometric wing characters to discriminate female Culex pipiens and Culex torrentium. J. Vector Ecol. 2014;39:204-212.

    Not in our library10.1111/j.1948-7134.2014.12088.x

  100. 100.

    Vogels C.B.F., van de Peppel L.J.J., van Vliet A.J.H., Westenberg M., Ibañez-Justicia A., Stroo A., Buijs J.A., Visser T.M., Koenraadt C.J.M. Winter activity and aboveground hybridization between the two biotypes of the West Nile virus vector Culex pipiens. Vector-Borne Zoonotic Dis. 2015;15:619-626.

    Not in our library10.1089/vbz.2015.1820

  101. 101.

    Danabalan R., Ponsonby D.J., Linton Y.-M. A critical assessment of available molecular identification tools for determining the status of Culex pipiens s.L. in the United Kingdom. J. Am. Mosq. Control Assoc. 2012;28:68-74.

    Not in our library10.2987/8756-971x-28.0.68

  102. 102.

    Shaikevich E.V., Vinogradova E.B., Bouattour A., Gouveia de Almeida A.P. Genetic diversity of Culex pipiens mosquitoes in distinct populations from Europe: Contribution of Cx. quinquefasciatus in Mediterranean populations. Parasit. Vectors. 2016;9:47.

    Not in our library10.1186/s13071-016-1333-8

  103. 103.

    Manley R., Harrup L.E., Veronesi E., Stubbins F., Stoner J., Gubbins S., Wilson A., Batten C., Koenraadt C.J.M., Henstock M., et al. Testing of UK populations of Culex pipiens L. for Schmallenberg virus vector competence and their colonization. PLoS ONE. 2015;10:e0134453.

    Not in our library10.1371/journal.pone.0134453

  104. 104.
  105. 105.

    Lunt D.H., Zhang D.-X., Szymura J.M., Hewitt O.M. The insect cytochrome oxidase I gene: Evolutionary patterns and conserved primers for phylogenetic studies. Insect Mol. Biol. 1996;5:153-165.

    Not in our library10.1111/j.1365-2583.1996.tb00049.x

  106. 106.

    Bernasconi M.V., Valsangiacomo C., Piffaretti J.C., Ward P.I. Phylogenetic relationships among Muscoidea (Diptera: Calyptratae) based on mitochondrial DNA sequences. Insect Mol. Biol. 2000;9:67-74.

    Not in our library10.1046/j.1365-2583.2000.00158.x

  107. 107.

    Shaikevich E.V., Zakharov I.A. Polymorphism of mitochondrial COI and nuclear ribosomal ITS2 in the Culex pipiens complex and in Culex torrentium (Diptera: Culicidae). Comp. Cytogenet. 2010;4:161-174.

    Not in our library10.3897/compcytogen.v4i2.45

  108. 108.

    Bourguet D., Fonseca D., Vourch G., Dubois M.P., Chandre F., Severini C., Raymond M. The acetylcholinesterase gene Ace: A diagnostic marker for the Pipiens and Quinquefasciatus forms of the Culex pipiens complex. J. Am. Mosq. Control Assoc. 1998;14:390-396.

    Not in our library

  109. 109.

    Atyame C.M., Delsuc F., Pasteur N., Weill M., Duron O. Diversification of Wolbachia endosymbiont in the Culex pipiens mosquito. Mol. Biol. Evol. 2011;28:2761-2772.

    Not in our library10.1093/molbev/msr083

  110. 110.

    Rudolf M., Czajka C., Börstler J., Melaun C., Jöst H., von Thien H., Badusche M., Becker N., Schmidt-Chanasit J., Krüger A., et al. First nationwide surveillance of Culex pipiens complex and Culex torrentium mosquitoes demonstrated the presence of Culex pipiens biotype pipiens/molestus hybrids in Germany. PLoS ONE. 2013;8:e71832.

    Not in our library10.1371/journal.pone.0071832

  111. 111.

    Yssouf A., Parola P., Lindström A., Lilja T., L’Ambert G., Bondesson U., Berenger J.M., Raoult D., Almeras L. Identification of European mosquito species by MALDI-TOF MS. Parasitol. Res. 2014;113:2375-2378.

    Not in our library10.1007/s00436-014-3876-y

  112. 112.

    Kaufmann C., Schaffner F., Ziegler D., Pflüger V., Mathis A. Identification of field-caught Culicoides biting midges using matrix-assisted laser desorption/ionization time of flight mass spectrometry. Parasitology. 2012;139:248-258.

    Not in our library10.1017/s0031182011001764

  113. 113.

    Yssouf A., Flaudrops C., Drali R., Kernif T., Socolovschi C., Berenger J.M., Raoult D., Parola P. Matrix-assisted laser desorption ionization-time of flight mass spectrometry for rapid identification of tick vectors. J. Clin. Microbiol. 2013;51:522-528.

    Not in our library10.1128/jcm.02665-12

  114. 114.

    Zittra C., Flechl E., Kothmayer M., Vitecek S., Rossiter H., Zechmeister T., Fuehrer H.-P. Ecological characterization and molecular differentiation of Culex pipiens complex taxa and Culex torrentium in eastern Austria. Parasit. Vectors. 2016;9:197.

    Not in our library10.1186/s13071-016-1495-4

  115. 115.

    Reusken C.B.E.M., De Vries A., Buijs J., Braks M.A.H., Den Hartog W., Scholte E.J. First evidence for presence of Culex pipiens biotype molestus in the Netherlands, and of hybrid biotype pipiens and molestus in Northern Europe. J. Vector Ecol. 2010;35:210-212.

    Not in our library10.1111/j.1948-7134.2010.00080.x

  116. 116.

    Bravo-Barriga D., Gomes B., Almeida A.P.G., Serrano-Aguilera F.J., Pérez-Martín J.E., Calero-Bernal R., Reina D., Frontera E., Pinto J. The mosquito fauna of the western region of Spain with emphasis on ecological factors and the characterization of Culex pipiens forms. J. Vector Ecol. 2017;42:136-147.

    Not in our library10.1111/jvec.12248

  117. 117.

    Osório H.C., Zé-Zé L., Amaro F., Nunes A., Alves M.J. Sympatric occurrence of Culex pipiens (Diptera, Culicidae) biotypes pipiens, molestus and their hybrids in Portugal, Western Europe: Feeding patterns and habitat determinants. Med. Vet. Entomol. 2014;28:103-109.

    Not in our library10.1111/mve.12020

  118. 118.
  119. 119.
  120. 120.

    Vogels C.B.F., Möhlmann T.W.R., Melsen D., Favia G., Wennergren U., Koenraadt C.J.M. Latitudinal diversity of Culex pipiens biotypes and hybrids in farm, peri-urban, and wetland habitats in Europe. PLoS ONE. 2016;11:e0166959.

    Not in our library10.1371/journal.pone.0166959

  121. 121.

    Shaikevich E.V., Vinogradova E.B. The discovery of a hybrid population of mosquitoes of the Culex pipiens L. complex (Diptera, Culicidae) on the Kos Island (Greece) by means of molecular markers. Entomol. Rev. 2014;94:35-39.

    Not in our library10.1134/s0013873814010047

  122. 122.

    Amraoui F., Tijane M., Sarih M., Failloux A.-B. Molecular evidence of Culex pipiens form molestus and hybrids pipiens/molestus in Morocco, North Africa. Parasit. Vectors. 2012;5:83.

    Not in our library10.1186/1756-3305-5-83

  123. 123.

    Cornel A.J., McAbee R.D., Rasgon J., Stanich M.A., Scott T.W., Coetzee M. Differences in extent of genetic introgression between sympatric Culex pipiens and Culex quinquefasciatus (Diptera: Culicidae) in California and South Africa. J. Med. Entomol. 2003;40:36-51.

    Not in our library10.1603/0022-2585-40.1.36

  124. 124.

    Vogels C.B.F., Hartemink N., Koenraadt C.J.M. Modelling West Nile virus transmission risk in Europe: Effect of temperature and mosquito biotypes on the basic reproduction number. Sci. Rep. 2017;7:5022.

    Not in our library10.1038/s41598-017-05185-4

  125. 125.

    Han L.L., Popovici F., Alexander J.P., Laurentia V., Tengelsen L.A., Cernescu C., Gary H.E., Ion-Nedelcu N., Campbell G.L., Tsai T.F., et al. Risk factors for West Nile virus infection and meningoencephalitis, Romania, 1996. J. Infect. Dis. 1999;179:230-233.

    Not in our library10.1086/314566

  126. 126.

    Fritz M.L., Walker E.D., Miller J.R., Severson D.W., Dworkin I. Divergent host preferences of above- and below-ground Culex pipiens mosquitoes and their hybrid offspring. Med. Vet. Entomol. 2015;29:115-123.

    Not in our library10.1111/mve.12096

  127. 127.
  128. 128.

    Weitzel T., Jawień P., Rydzanicz K., Lonc E., Becker N. Culex pipiens s.l. and Culex torrentium (Culicidae) in Wrocław area (Poland): Occurrence and breeding site preferences of mosquito vectors. Parasitol. Res. 2015;114:289-295.

    Not in our library10.1007/s00436-014-4193-1

  129. 129.

    Lühken R., Steinke S., Leggewie M., Tannich E., Krüger A., Becker S., Kiel E. Physico-chemical characteristics of Culex pipiens sensu lato and Culex torrentium (Diptera: Culicidae) breeding sites in Germany. J. Med. Entomol. 2015;52:932-936.

    Not in our library10.1093/jme/tjv070

  130. 130.

    Service M.W. The taxonomy and biology of two sympatric sibling species of Culex, C. pipiens and C. torrentium (Diptera, Culicidae). J. Zool. 1968;156:313-323.

    Not in our library10.1111/j.1469-7998.1968.tb04355.x

  131. 131.

    Simpson J.E., Hurtado P.J., Medlock J., Molaei G., Andreadis T.G., Galvani A.P., Diuk-Wasser M.A. Vector host-feeding preferences drive transmission of multi-host pathogens: West Nile virus as a model system. Proc. R. Soc. B Biol. Sci. 2012;279:925-933.

    Not in our library10.1098/rspb.2011.1282

  132. 132.

    Takken W., Verhulst N.O. Host preferences of blood-feeding mosquitoes. Annu. Rev. Entomol. 2013;58:433-453.

    Not in our library10.1146/annurev-ento-120811-153618

  133. 133.
  134. 134.

    Edman J.D., Scott T.W. Host defensive behaviour and the feeding success of mosquitoes. Int. J. Trop. Insect Sci. 1987;8:617-622.

    Not in our library10.1017/s1742758400022694

  135. 135.

    De Boer J.G., Robinson A., Powers S.J., Burgers S.L.G.E., Caulfield J.C., Birkett M.A., Smallegange R.C., van Genderen P.J.J., Bousema T., Sauerwein R.W., et al. Odours of Plasmodium falciparum-infected participants influence mosquito-host interactions. Sci. Rep. 2017;7:9283.

    Not in our library10.1038/s41598-017-08978-9

  136. 136.

    Cornet S., Nicot A., Rivero A., Gandon S. Both infected and uninfected mosquitoes are attracted toward malaria infected birds. Malar. J. 2013;12:179.

    Not in our library10.1186/1475-2875-12-179

  137. 137.

    Cornet S., Nicot A., Rivero A., Gandon S. Malaria infection increases bird attractiveness to uninfected mosquitoes. Ecol. Lett. 2013;16:323-329.

    Not in our library10.1111/ele.12041

  138. 138.

    Vogels C.B.F., Fros J.J., Pijlman G.P., van Loon J.J.A., Gort G., Koenraadt C.J.M. Virus interferes with host-seeking behaviour of mosquito. J. Exp. Biol. 2017;220:3598-3603.

    Not in our library10.1242/jeb.164186

  139. 139.

    Busula A.O., Bousema T., Mweresa C.K., Masiga D., Logan J.G., Sauerwein R.W., Verhulst N.O., Takken W., de Boer J.G. Gametocytemia and attractiveness of Plasmodium falciparum-infected Kenyan children to Anopheles gambiae mosquitoes. J. Infect. Dis. 2017;216:291-295.

    Not in our library10.1093/infdis/jix214

  140. 140.

    Harbach R.E., Harrison B.A., Gad A.M. Culex (Culex) molestus Forskal (Diptera: Culicidae): Neotype designation, description, variation and taxonomic status. Proc. Entomol. Soc. Washingt. 1984;86:521-542.

    Not in our library

  141. 141.

    Harbach R.E., Dahl C., White G.B. Culex (Culex) pipiens Linnacus (Diptera: Culicidae): Concepts, type designations, and description. Proc. Entomol. Soc. Washingt. 1985;87:1-24.

    Not in our library

  142. 142.

    Platonov A.E., Fedorova M.V., Karan L.S., Shopenskaya T.A., Platonova O.V., Zhuravlev V.I. Epidemiology of West Nile infection in Volgograd, Russia, in relation to climate change and mosquito (Diptera: Culicidae) bionomics. Parasitol. Res. 2008;103.

    Not in our library10.1007/s00436-008-1050-0

  143. 143.

    Martínez-de la Puente J., Moreno-Indias I., Hernández-Castellano L.E., Argüello A., Ruiz S., Soriguer R., Figuerola J. Host-feeding pattern of Culex theileri (Diptera: Culicidae), potential vector of Dirofilaria immitis in the Canary Islands, Spain. J. Med. Entomol. 2012;49:1419-1423.

    Not in our library10.1603/me12081

  144. 144.

    Osório H.C., Zé-Zé L., Alves M.J. Host-feeding patterns of Culex pipiens and other potential mosquito vectors (Diptera: Culicidae) of West Nile virus (Flaviviridae) collected in Portugal. J. Med. Entomol. 2012;49:717-721.

    Not in our library10.1603/me11184

  145. 145.

    Korkmaz S., Yildirim A., Duzlu O., Ciloglu A., Onder Z., Inci A. Blood meal identification of the mosquito (Diptera: Culicidae) specimens belong to Culex pipiens complex that were collected from Kayseri Province. Turkish J. Parasitol. 2017;40:199-204.

    Not in our library10.5152/tpd.2016.4882

  146. 146.

    Martínez-de la Puente J., Muñoz J., Capelli G., Montarsi F., Soriguer R., Arnoldi D., Rizzoli A., Figuerola J. Avian malaria parasites in the last supper: identifying encounters between parasites and the invasive Asian mosquito tiger and native mosquito species in Italy. Malar. J. 2015;14:32.

    Not in our library10.1186/s12936-015-0571-0

  147. 147.

    Martínez-de la Puente J., Ferraguti M., Ruiz S., Roiz D., Soriguer R.C., Figuerola J. Culex pipiens forms and urbanization: Effects on blood feeding sources and transmission of avian Plasmodium. Malar. J. 2016;15:589.

    Not in our library10.1186/s12936-016-1643-5

  148. 148.

    Börstler J., Jöst H., Garms R., Krüger A., Tannich E., Becker N., Schmidt-Chanasit J., Lühken R. Host-feeding patterns of mosquito species in Germany. Parasit. Vectors. 2016;9:318.

    Not in our library10.1186/s13071-016-1597-z

  149. 149.

    Gomes B., Sousa C.A., Vicente J.L., Pinho L., Calderón I., Arez E., Almeida A.P.G., Donnelly M.J., Pinto J. Feeding patterns of molestus and pipiens forms of Culex pipiens (Diptera: Culicidae) in a region of high hybridization. Parasit. Vectors. 2013;6:93.

    Not in our library10.1186/1756-3305-6-93

  150. 150.
  151. 151.

    Martínez-de la Puente J., Ruiz S., Soriguer R., Figuerola J. Effect of blood meal digestion and DNA extraction protocol on the success of blood meal source determination in the malaria vector Anopheles atroparvus. Malar. J. 2013;12:109.

    Not in our library10.1186/1475-2875-12-109

  152. 152.

    Muñoz J., Eritja R., Alcaide M., Montalvo T., Soriguer R.C., Figuerola J. Host-feeding patterns of native Culex pipiens and invasive Aedes albopictus mosquitoes (Diptera: Culicidae) in urban zones from Barcelona, Spain. J. Med. Entomol. 2011;48:956-960.

    Not in our library10.1603/me11016

  153. 153.

    Cancrini G., Magi M., Gabrielli S., Arispici M., Tolari F., Dell’Omodarme M., Prati M.C. Natural vectors of Dirofilariasis in rural and urban areas of the Tuscan region, central Italy. J. Med. Entomol. 2006;43:574-579.

    Not in our library10.1093/jmedent/43.3.574

  154. 154.

    Curtotti A. Characterization of East London Culex pipiens s.l. in Relation to the Risk of Transmission to Humans of the West Nile Virus in Great Britain. PhD Thesis. 2009.

    Not in our library

  155. 155.

    Radrova J., Seblova V., Votypka J. Feeding behavior and spatial distribution of Culex mosquitoes (Diptera: Culicidae) in wetland areas of the Czech Republic. J. Med. Entomol. 2013;50:1097-1104.

    Not in our library10.1603/me13029

  156. 156.

    Schönenberger A.C., Wagner S., Tuten H.C., Schaffner F., Torgerson P., Furrer S., Mathis A., Silaghi C. Host preferences in host-seeking and blood-fed mosquitoes in Switzerland. Med. Vet. Entomol. 2016;30:39-52.

    Not in our library10.1111/mve.12155

  157. 157.

    Service M.W. Feeding behaviour and host preferences of British mosquitoes. Bull. Entomol. Res. 1971;60:653-661.

    Not in our library10.1017/s0007485300042401

  158. 158.

    Service M.W. A reappraisal of the role of mosquitoes in the transmission of myxomatosis in Britain. J. Hyg. (Lond). 1971;69:105-111.

    Not in our library10.1017/s0022172400021318

  159. 159.

    Balenghien T., Fouque F., Sabatier P., Bicout D.J. Horse-, bird-, and human-seeking behavior and seasonal abundance of mosquitoes in a West Nile virus focus of southern France. J. Med. Entomol. 2006;43:936-946.

    Not in our library10.1093/jmedent/43.5.936

  160. 160.

    Service M.W. Observations on the ecology of some British mosquitoes. Bull. Entomol. Res. 1969;59:161-193.

    Not in our library10.1017/s000748530000314x

  161. 161.

    Rizzoli A., Bolzoni L., Chadwick E.A., Capelli G., Montarsi F., Grisenti M., Martínez-de la Puente J., Muñoz J., Figuerola J., Soriguer R., et al. Understanding West Nile virus ecology in Europe: Culex pipiens host feeding preference in a hotspot of virus emergence. Parasit. Vectors. 2015;8:213.

    Not in our library10.1186/s13071-015-0831-4

  162. 162.

    Fyodorova M.V., Savage H.M., Lopatina J.V., Bulgakova T.A., Ivanitsky A.V., Platonova O.V., Platonov A.E. Evaluation of potential West Nile virus vectors in Volgograd Region, Russia, 2003 (Diptera: Culicidae): Species composition, bloodmeal host utilization, and virus infection rates of mosquitoes. J. Med. Entomol. 2006;43:552-563.

    Not in our library10.1093/jmedent/43.3.552

  163. 163.

    Jaenson T.G.T., Niklasson B. Feeding patterns of mosquitoes (Diptera: Culicidae) in relation to the transmission of Ockelbo disease in Sweden. Bull. Entomol. Res. 1986;76:375.

    Not in our library10.1017/s0007485300014863

  164. 164.

    Muñoz J., Ruiz S., Soriguer R., Alcaide M., Viana D.S., Roiz D., Vázquez A., Figuerola J. Feeding patterns of potential West Nile virus vectors in south-west Spain. PLoS ONE. 2012;7:e39549.

    Not in our library10.1371/journal.pone.0039549

  165. 165.

    Onyeka J.O.A., Boreham P.F.L. Population studies, physiological state and mortality factors of overwintering adult populations of females of Culex pipiens L. (Diptera:Culicidae). Bull. Entomol. Res. 1987;77:99-112.

    Not in our library10.1017/s0007485300011585

  166. 166.

    Ferraguti M., Martínez-de la Puente J., Muñoz J., Roiz D., Ruiz S., Soriguer R., Figuerola J. Avian Plasmodium in Culex and Ochlerotatus mosquitoes from Southern Spain: Effects of season and host-feeding source on parasite dynamics. PLoS ONE. 2013;8:e66237.

    Not in our library10.1371/journal.pone.0066237

  167. 167.

    Roiz D., Vazquez A., Rosà R., Muñoz J., Arnoldi D., Rosso F., Figuerola J., Tenorio A., Rizzoli A. Blood meal analysis, flavivirus screening, and influence of meteorological variables on the dynamics of potential mosquito vectors of West Nile virus in northern Italy. J. Vector Ecol. 2012;37:20-28.

    Not in our library10.1111/j.1948-7134.2012.00196.x

  168. 168.

    Brugman V.A., Hernández-Triana L.M., England M.E., Medlock J.M., Mertens P.P.C., Logan J.G., Wilson A.J., Fooks A.R., Johnson N., Carpenter S. Blood-feeding patterns of native mosquitoes and insights into their potential role as pathogen vectors in the Thames estuary region of the United Kingdom. Parasites Vectors. 2017;10:163.

    Not in our library10.1186/s13071-017-2098-4

  169. 169.

    Hernández-Triana L.M., Brugman V.A., Prosser S.W.J., Weland C., Nikolova N., Thorne L., Fernández de Marco M., Fooks A.R., Johnson N. Molecular approaches for blood meal analysis and species identification of mosquitoes (Insecta: Diptera: Culicidae) in rural locations in southern England, United Kingdom. Zootaxa. 2017;4250:67.

    Not in our library10.11646/zootaxa.4250.1.5

  170. 170.

    Kilpatrick A.M., Kramer L.D., Jones M.J., Marra P.P., Daszak P., Fonseca D.M. Genetic influences on mosquito feeding behavior and the emergence of zoonotic pathogens. Am. J. Trop. Med. Hyg. 2007;77:667-671.

    Not in our library

  171. 171.

    Huang S., Hamer G.L., Molaei G., Walker E.D., Goldberg T.L., Kitron U.D., Andreadis T.G. Genetic variation associated with mammalian feeding in Culex pipiens from a West Nile virus epidemic region in Chicago, Illinois. Vector-Borne Zoonotic Dis. 2009;9:637-642.

    Not in our library10.1089/vbz.2008.0146

  172. 172.

    Silver J.B. Mosquito Ecology: Field Sampling Methods. 2007.

    Not in our library

  173. 173.
  174. 174.

    Sieg M., Schmidt V., Ziegler U., Keller M., Höper D., Heenemann K., Rückner A., Nieper H., Muluneh A., Groschup M.H., et al. Outbreak and Cocirculation of Three Different Usutu Virus Strains in Eastern Germany. Vector-Borne Zoonotic Dis. 2017;17:662-664.

    Not in our library10.1089/vbz.2016.2096

  175. 175.

    Medlock J.M., Vaux A.G.C. Impacts of the creation, expansion and management of English wetlands on mosquito presence and abundance—Developing strategies for future disease mitigation. Parasit. Vectors. 2015;8:142.

    Not in our library10.1186/s13071-015-0751-3

  176. 176.

    Townroe S., Callaghan A. British container breeding mosquitoes: The impact of urbanisation and climate change on community composition and phenology. PLoS ONE. 2014;9:e95325.

    Not in our library10.1371/journal.pone.0095325

  177. 177.

    Dekoninck W., Hendrickx F., Vasn Bortel W., Versteirt V., Coosemans M., Damiens D., Hance T., De Clercq E.M., Hendrickx G., Schaffner F., et al. Human-induced expanded distribution of Anopheles plumbeus, experimental vector of West Nile virus and a potential vector of human malaria in Belgium. J. Med. Entomol. 2011;48:924-928.

    Not in our library10.1603/me10235

  178. 178.

    Medlock J.M., Vaux A.G.C. Colonization of a newly constructed urban wetland by mosquitoes in England: Implications for nuisance and vector species. J. Vector Ecol. 2014;39:249-260.

    Not in our library10.1111/jvec.12099

  179. 179.

    Kassim N.F.A., Webb C.E., Russell R.C. Is the expression of autogeny by Culex molestus Forskal (Diptera: Culicidae) influenced by larval nutrition or by adult mating, sugar feeding, or blood feeding? J. Vector Ecol. 2012;37:162-171.

    Not in our library10.1111/j.1948-7134.2012.00213.x

  180. 180.

    Rudolf I., Betášová L., Blažejová H., Venclíková K., Straková P., Šebesta O., Mendel J., Bakonyi T., Schaffner F., Nowotny N., et al. West Nile virus in overwintering mosquitoes, central Europe. Parasit. Vectors. 2017;10:452.

    Not in our library10.1186/s13071-017-2399-7

  181. 181.

    Ciota A.T., Drummond C.L., Drobnack J., Ruby M.A., Kramer L.D., Ebel G.D. Emergence of Culex pipiens from overwintering hibernacula. J. Am. Mosq. Control Assoc. 2011;27:21-29.

    Not in our library10.2987/8756-971x-27.1.21

  182. 182.

    Nelms B.M., Macedo P.A., Kothera L., Savage H.M., Reisen W.K. Overwintering biology of Culex (Diptera: Culicidae) mosquitoes in the Sacramento Valley of California. J. Med. Entomol. 2013;50:773-790.

    Not in our library10.1603/me12280

  183. 183.

    Roiz D., Ruiz S., Soriguer R., Figuerola J. Climatic effects on mosquito abundance in Mediterranean wetlands. Parasit. Vectors. 2014;7:333.

    Not in our library10.1186/1756-3305-7-333

  184. 184.
  185. 185.

    LaDeau S.L., Allan B.F., Leisnham P.T., Levy M.Z. The ecological foundations of transmission potential and vector-borne disease in urban landscapes. Funct. Ecol. 2015;29:889-901.

    Not in our library10.1111/1365-2435.12487

  186. 186.

    Ibañez-Justicia A., Cianci D. Modelling the spatial distribution of the nuisance mosquito species Anopheles plumbeus (Diptera: Culicidae) in the Netherlands. Parasit. Vectors. 2015;8:258.

    Not in our library10.1186/s13071-015-0865-7

  187. 187.

    Heym E.C., Kampen H., Fahle M., Hohenbrink T.L., Schäfer M., Scheuch D.E., Walther D. Anopheles plumbeus (Diptera: Culicidae) in Germany: updated geographic distribution and public health impact of a nuisance and vector mosquito. Trop. Med. Int. Heal. 2017;22:103-112.

    Not in our library10.1111/tmi.12805

  188. 188.

    Misslin R., Telle O., Daudé E., Vaguet A., Paul R.E. Urban climate versus global climate change-what makes the difference for dengue? Ann. N. Y. Acad. Sci. 2016;1382:56-72.

    Not in our library10.1111/nyas.13084

  189. 189.

    Brown L., Medlock J.M., Murray V. Impact of drought on vector-borne diseases—how does one manage the risk? Public Health. 2014;128:29-37.

    Not in our library10.1016/j.puhe.2013.09.006

  190. 190.

    Beji M., Rhim A., Roiz D., Bouattour A. Ecophysiological characterization and molecular differentiation of Culex pipiens forms (Diptera: Culicidae) in Tunisia. Parasit. Vectors. 2017;10:327.

    Not in our library10.1186/s13071-017-2265-7

  191. 191.

    Gibson G., Russell I. Flying in tune: Sexual recognition in mosquitoes. Curr. Biol. 2006;16:1311-1316.

    Not in our library10.1016/j.cub.2006.05.053

  192. 192.

    Pennetier C., Warren B., Dabiré K.R., Russell I.J., Gibson G. “Singing on the Wing” as a mechanism for species recognition in the malarial mosquito Anopheles gambiae. Curr. Biol. 2010;20:131-136.

    Not in our library10.1016/j.cub.2009.11.040

  193. 193.

    Duron O., Raymond M., Weill M. Many compatible Wolbachia strains coexist within natural populations of Culex pipiens mosquito. Heredity (Edinb.). 2011;106:986-993.

    Not in our library10.1038/hdy.2010.146

  194. 194.

    Sanders C.J., Harrup L.E., Tugwell L.A., Brugman V.A., England M., Carpenter S. Quantification of within- and between-farm dispersal of Culicoides biting midges using an immunomarking technique. J. Appl. Ecol. 2017;54:1429-1439.

    Not in our library10.1111/1365-2664.12875

  195. 195.

    Verhulst N.O., Loonen J.A.C.M., Takken W. Advances in methods for colour marking of mosquitoes. Parasit. Vectors. 2013;6:200.

    Not in our library10.1186/1756-3305-6-200

  196. 196.

    Syed Z., Leal W.S. Acute olfactory response of Culex mosquitoes to a human- and bird-derived attractant. Proc. Natl. Acad. Sci. USA. 2009;106:18803-18808.

    Not in our library10.1073/pnas.0906932106

  197. 197.

    Allan S.A., Bernier U.R., Kline D.L. Laboratory evaluation of avian odors for mosquito (Diptera: Culicidae) attraction. J. Med. Entomol. 2006;43:225-231.

    Not in our library10.1093/jmedent/43.2.225

  198. 198.

    Mauer D.J., Rowley W.A. Attraction of Culex pipiens pipiens (Diptera: Culicidae) to flower volatiles. J. Med. Entomol. 1999;36:503-507.

    Not in our library10.1093/jmedent/36.4.503

  199. 199.

    Jhumur U.S., Dötterl S., Jürgens A. Naïve and conditioned responses of Culex pipiens pipiens biotype molestus (Diptera: Culicidae) to flower odors. J. Med. Entomol. 2006;43:1164-1170.

    Not in our library10.1093/jmedent/43.6.1164

  200. 200.

    Hesson J.C., Ignell R., Hill S.R., Östman Ö., Lundström J.O. Trapping biases of Culex torrentium and Culex pipiens revealed by comparison of captures in CDC traps, ovitraps, and gravid traps. J. Vector Ecol. 2015;40:158-163.

    Not in our library10.1111/jvec.12145

  201. 201.

    Weitzel T., Braun K., Collado A., Jost A., Becker N. Distribution and frequency of Culex pipiens and Culex torrentium (Culicidae) in Europe and diagnostic allozyme markers. Eur. Mosq. Bull. 2011;29:22-37.

    Not in our library

  202. 202.

    Calzolari M., Pautasso A., Montarsi F., Albieri A., Bellini R., Bonilauri P., Defilippo F., Lelli D., Moreno A., Chiari M., et al. West Nile virus surveillance in 2013 via mosquito screening in Northern Italy and the influence of weather on virus circulation. PLoS ONE. 2015;10:e014915.

    Not in our library10.1371/journal.pone.0140915

  203. 203.

    Vaux A.G.C., Gibson G., Hernández-Triana L.M., Cheke R.A., McCracken F., Jeffries C.L., Horton D.L., Springate S., Johnson N., Fooks A.R., et al. Enhanced West Nile virus surveillance in the North Kent marshes, UK. Parasit. Vectors. 2015;8:91.

    Not in our library10.1186/s13071-015-0705-9

  204. 204.

    Boukraa S., de La Grandiere M.A., Bawin T., Raharimalala F.N., Zimmer J.Y., Haubruge E., Thiry E., Francis F. Diversity and ecology survey of mosquitoes potential vectors in Belgian equestrian farms: A threat prevention of mosquito-borne equine arboviruses. Prev. Vet. Med. 2016;124:58-68.

    Not in our library10.1016/j.prevetmed.2015.12.013

  205. 205.

    Gomes B., Kioulos E., Papa A., Almeida A.P.G., Vontas J., Pinto J. Distribution and hybridization of Culex pipiens forms in Greece during the West Nile virus outbreak of 2010. Infect. Genet. Evol. 2013;16:218-225.

    Not in our library10.1016/j.meegid.2013.02.006

  206. 206.

    Petrić D., Petrović T., Hrnjaković Cvjetković I., Zgomba M., Milošević V., Lazić G., Ignjatović Ćupina A., Lupulović D., Lazić S., Dondur D., et al. West Nile virus “circulation” in Vojvodina, Serbia: Mosquito, bird, horse and human surveillance. Mol. Cell. Probes. 2017;31:28-36.

    Not in our library10.1016/j.mcp.2016.10.011

  207. 207.

    Brustolin M., Talavera S., Santamaría C., Rivas R., Pujol N., Aranda C., Marquès E., Valle M., Verdún M., Pagès N., et al. Culex pipiens and Stegomyia albopicta (=Aedes albopictus) populations as vectors for lineage 1 and 2 West Nile virus in Europe. Med. Vet. Entomol. 2016;30:166-173.

    Not in our library10.1111/mve.12164

  208. 208.

    Rogozi E., Velo E., Bino S., Kadriaj P., Severini F., Schaffner F. An update of the mosquito fauna of Albania, based on a country-wide field survey 2011–2012. Proceedings of the E-Sove: From biology to integrated in a changing world.

    Not in our library

  209. 209.

    Snow K., Ramsdale C. Distribution chart for European mosquitoes. Eur. Mosq. Bull. 1999;3:14-31.

    Not in our library

  210. 210.

    Boukraa S., Dekoninck W., Versteirt V., Schaffner F., Coosemans M., Haubruge E., Francis F. Updated checklist of the mosquitoes (Diptera: Culicidae) of Belgium. J. Vector Ecol. 2015;40:398-407.

    Not in our library10.1111/jvec.12180

  211. 211.

    Huldén L., Huldén L. Checklist of the family Culicidae (Diptera) in Finland. Zookeys. 2014.

    Not in our library10.3897/zookeys.441.7743

  212. 212.

    Culverwell C.L. A report on the mosquitoes of mainland Åland, southwestern Finland and revised list of Finnish mosquitoes. Med. Vet. Entomol. 2017.

    Not in our library10.1111/mve.12272

  213. 213.

    Schaffner F. A revised cheddist of the French Culicidae. Eur. Mosq. Bull. 1998;2:1-9.

    Not in our library

  214. 214.

    Tóth S., Kenyeres Z. Revised checklist and distribution maps of mosquitoes (Diptera, Culicidae) of Hungary. Eur. Mosq. Bull. 2012;30:30-65.

    Not in our library

  215. 215.

    Pakalniškis S., Bernotienė R., Lutovinovas E., Petrašiūnas A., Podėnas S., Rimšaitė J., Saether O.A., Spungis V. Checklist of Lithuanian Diptera. New Rare Lith. Insect Species. 2006;18:16-154.

    Not in our library10.1080/13921657.2000.10512316

  216. 216.

    Beck M., Galm M., Weitzel T., Fohlmeister V., Kaiser A., Amold A. Preliminary studies on the mosquito fauna of Luxembourg. Eur. Mosq. Bull. 2003;14:21-24.

    Not in our library

  217. 217.

    Sulesco T.M., Toderas I.K., Toderas L.G. Annotated checklist of the mosquitoes of the Republic of Moldova. J. Am. Mosq. Control Assoc. 2013;29:98-101.

    Not in our library10.2987/12-6311r.1

  218. 218.

    Ribeiro H., Ramos H., Pires C., Capela R. An annotated checklist of the mosquitoes of continental Portugal (Diptera, Culicidae). Acta III Congr. Iber. Entomol. Spec. 1988;233-254.

    Not in our library

  219. 219.

    Nicolescu G., Vladimirescu A., Ciolpan O. The distribution of mosquitoes in Romania (Diptera: Culicidae). Part I: Anopheles, Aedes and Culex. Eur. Mosq. Bull. 2002;13:17-26.

    Not in our library

  220. 220.

    Božičić-Lothrop B., Vujić A. Fauna of mosquitoes (Diptera: Culicidae) of Stara Planina, Serbia. Acta Entomol. Serbica. 1996;1:31-38.

    Not in our library

  221. 221.

    Jalili N., Országh I., Halgoš J., Labuda M. Mosquitoes (Diptera, Culicidae) of Slovakia. Eur. Mosq. Bull. 2000;6:20-26.

    Not in our library

  222. 222.

    Aranda C., Eritja R., Schaffner F., Escosa R. Culex (Culex) torrentium Martini (Diptera: Culicidae) a new species from Spain. Eur. Mosq. Bull. 2000;8:7-9.

    Not in our library

  223. 223.

    Eritja R., Aranda C., Padrós J., Goula M., Lucientes J., Escosa R., Marques E., Cáceres F. An annotaded checklist and bibliography of the mosquitoes of Spain (Diptera: Culicidae). Eur. Mosq. Bull. 2000;8:10-18.

    Not in our library10.2987/8756-971x(2008)24[i:jotamc]2.0.co;2

  224. 224.

    Gunay F., Alten B., Simsek F., Aldemir A., Linton Y.M. Barcoding Turkish Culex mosquitoes to facilitate arbovirus vector incrimination studies reveals hidden diversity and new potential vectors. Acta Trop. 2015;143:112-120.

    Not in our library10.1016/j.actatropica.2014.10.013

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