INTRODUCTION
Mosquitoes (Diptera: Culicidae) are the world’s deadliest animals,1 vectoring myriad pathogens that result in untold pain and misery. Seventy nine native mosquito species are known from Canada,2–4 though several more are suspected to occur based on their distributions in the United States,4 and five invasive species are currently known.2,4,5 In this review, we define invasive mosquitoes as those that have been accidentally or deliberately introduced into areas beyond their native range and whose presence negatively impact the environment, the economy, or the society including human and animal health.6 Invasive mosquitoes threaten human and animal health as they can vector pathogens not previously known from Canada, or act as more efficient vectors of native pathogens. While the permanent establishment of many exotic mosquito-borne pathogens discussed here is unlikely in Canada, this may not always be the case due to novel mutations or climate change. Furthermore, localized seasonal outbreaks resulting from travel-related or other imported cases have occurred elsewhere.7 With increasing globalization and the landscape of emerging pathogens constantly changing, and as demonstrated by literature for other areas,8 an entomological, medical, and veterinary knowledge of the invasive mosquitoes of Canada is more important than ever.
IMPORTANT MOSQUITO-BORNE PATHOGENS CURRENTLY OR FORMERLY ENDEMIC TO CANADA
Snowshoe hare virus (SSHV; Family: Bunyaviridae, genus: Bunyavirus) undergoes an enzootic transmission cycle in wild mammals with mosquitoes with non-Culex mosquitoes acting as the primary vectors.9 It is not clear what species are the principal hosts of SSHV, although small mammals are thought to be important in SSHV maintenance and amplification.9 Snowshoe hare virus is found across Canada10 as well as Alaska11 and parts of northern Eurasia.12 This virus has been reported to cause clinical encephalitis in humans,13 predominantly in children,9 and horses.14
Dog heartworm, Dirofilaria immitis, is a parasitic filarial worm that is an obligate parasite of mosquitoes and canids,15 although rare cases in other animals, such as humans, also occur.16 Endemic foci of D. immitis occur in many parts of Canada, particularly southern regions.17,18 A variety of mosquito species in the genera Aedes, Culex, and Anopheles vector D. immitis.19
West Nile virus (WNV; Family: Flaviviridae, genus: Flavivirus) undergoes an enzootic transmission cycle in avian hosts with mosquitoes in the genus Culex acting as primary vectors,20,21 though other mosquitoes with wide host ranges can carry WNV as well.22 The primary vectors of WNV in Canada are Culex pipiens and Cx. restuans in Eastern Canada, and Cx. tarsalis in Western Canada.20–25 Culex spp. are often ornithophilic, with some species feeding on humans as well. Ornithophilic biting behavior in the spring, by early emerging Culex spp. (such as Cx. restuans in Ontario) may vector enzootic transmission within local or migratory bird populations.26 During late summer vector species such as Cx. tarsalis or Cx. pipiens may increase their biting of humans27 leading to the transmission of WNV from birds to humans. West Nile virus first arrived in North America in 1999,28 and was first detected in Canada during 2001 in Ontario.29 By 2009, it had spread all the way west to British Columbia.25 West Nile virus can cause disease in several animals, including mortality in horses and birds.30,31
St. Louis encephalitis virus (SLEV; Family: Flaviviridae, genus: Flavivirus) undergoes an enzootic transmission cycle among birds and is vectored mainly by mosquitoes of the genus Culex.32 It may undergo vertical transmission and persist in mosquitoes through winter.32,33 During the 1970s, Canada experienced epidemics of SLEV, with the virus reported in Saskatchewan, Manitoba, Ontario, and Quebec,10 although in the United States it has been reported from coast-to-coast but predominantly in the southern States.32 St. Louis encephalitis virus can cause disease in horses as well.34
Jamestown Canyon virus (JCV; Family: Peribunyaviridae, genus: Orthobunyavirus) is transmitted primarily among wild ungulates by non-Culex mosquitoes.9 One study in the eastern United States found more than 20 field-collected mosquito species tested positive for JCV, with Anopheles punctipennis, Coquillettidia perturbans, and several Aedes spp. incriminated as likely vectors.35 Jamestown Canyon virus is widespread in temperate North America and, while human infections and disease are rare, they are likely underrecognized.36
Cache Valley virus (CVV; Family: Peribunyaviridae, genus: Orthobunyavirus) is transmitted primarily among ungulates by non-Culex mosquitoes.9 In the Canadian prairies, CVV has been isolated from Aedes vexans, Culiseta incidens, and Culex tarsalis.37 In rare instances, CVV can cause disease in humans,38 and congenital malformations in sheep and goats.39,40
The human malaria parasite, Plasmodium vivax, is transmitted by mosquitoes of the genus Anophleles. Plasmodium vivax was formerly endemic to parts of Eastern Canada,41 and the southern interior of British Columbia.42 Contemporary imported cases continue to result in local malaria outbreaks.43
Eastern equine encephalitis virus (EEEV; Family: Togaviridae, genus: Alphavirus) undergoes an enzootic transmission cycle among passerine birds and mosquito vectors.32 Eastern equine encephalitis virus is vectored between birds by the mosquito Culiseta melanura,44 which does not bite humans.45 Transmission of EEEV to humans and other mammals occurs through mosquitoes that feed on passerine birds and mammals, including Coquillettidia perturbans, Cs. morsitans, Culex spp., and some mosquitoes of the genus Aedes.32,46 In Canada, EEEV is found in Ontario and Quebec and can cause mortality in humans and horses.44 Domestic poultry has been reported in some cases to suffer a decrease in egg production as a result of infection with EEEV,47 and even mortality.47
Western equine encephalitis virus (WEEV; Family: Togaviridae, genus: Alphavirus) is transmitted between birds and mammals by a variety of mosquitoes, although the western encephalitis mosquito, Cx. tarsalis, is thought to be the most important vector.32 Western equine encephalitis virus can cause mortality in humans and horses,32 and it may affect domestic poultry as well including decreased egg laying.48 In Canada, WEEV is found from British Columbia to the Great Lakes.10
SELECT EXOTIC PATHOGENS RELEVANT TO THIS REVIEW
Dengue virus (DENV; Family: Flaviviridae, genus: Flavivirus) is an arbovirus ubiquitous in the tropics that is vectored by some mosquitoes in the genus Aedes,49 primarily between humans but also nonhuman primates.50 There are several serotypes of dengue individuals who experience a subsequent infection with a different serotype are at increased risk of developing severe dengue.51
Japanese encephalitis (JEV; Family: Flaviviridae, genus: Flavivirus) undergoes enzootic transmission between birds and pigs and has recently spread from southeast Asia into Australia.52 This virus is the leading cause of encephalitis in eastern and southern Asia and is primarily vectored by mosquitoes of the genus Culex, although some Aedes spp. also act as vectors.52–54
Usutu virus (USUV; Family: Flaviviridae, genus: Flavivirus) is primarily vectored by Culex mosquitoes, and some members of the genus Aedes. Usutu virus was previously only known from Africa, but it has recently spread to Europe.55 Usutu virus primarily circulates in humans and birds, where it can cause encephalitis in humans56 and has caused severe mortality in bird populations that have not developed immunity.55
Yellow fever virus (YFV; Family: Flaviviridae, genus: Flavivirus) is transmitted among humans and other primates primarily by some Aedes, Sabethes, and Haemogogus mosquitoes.57 There is a vaccine for YFV; however, it has historically been considered a very dangerous pathogen.57 Yellow fever virus is primarily tropical in distribution; however, sporadic outbreaks have occurred as far north as New York City and Philadelphia,57 and there is risk of travel-related cases initiating autochthonous transmission cycles.58
Zika virus (ZIKV; Family: Flaviviridae, genus: Flavivirus) is transmitted between nonhuman primates and humans, primarily by many mosquitoes of the genus Aedes, although there are other transmission routes.59 Travellers returning from areas with endemic ZIKV may be at risk of initiating autochthonous transmission if competent vectors are present.59
Chikungunya (CHIKV; Family: Togaviridae, genus: Alphavirus) is primarily vectored by some Aedes mosquitoes.60 Nonhuman primates serve as potential reservoir or amplifying hosts,60 although vertical transmission in mosquitoes has also been reported.61
La Crosse virus (LACV; Family: Peribunyaviridae, genus: Orthobunyavirus) is the primary cause of viral encephalitis in children in the United States.9 The primary LACV vectors, Aedes triseriatus and Ae. albopictus, have a limited distribution within Canada.4,5
Rift Valley fever (RVFV; Family: Phenuiviridae, genus: Phlebovirus) is known from Africa and Arabia where it causes morbidity and mortality in humans and ruminants and is vectored by several pathways, including mosquitoes of a variety of genera.62
INVASIVE MOSQUITOES KNOWN FROM CANADA
Aedes aegypti distribution.
The yellow fever mosquito, Aedes (Stegomyia) aegypti (L.), originated in sub-Saharan Africa where its sylvatic form can still be found today.63 As it adapted to a synanthropic lifestyle, this species managed to spread to tropical and subtropical areas around the globe via human-assisted dispersal, particularly in association with ship traffic connected with the slave trade.64 One of the most globally widespread species in tropical and subtropical environments, Ae. aegypti has been present in North America for centuries but is intolerant of temperate winters.65 While Ae. aegypti has historically been limited to areas with mean January temperatures above 10°C,65 there are sporadic northern populations that exist in areas where mean January temperatures get as low as about 2°C.66 Habitat models for Ae. aegypti do not predict suitable year-round climate conditions for this species in Canada now or in the near future.67,68
In 2016 and 2017, low numbers of Ae. aegypti were reported from Southern Ontario, representing the first records of this species in Canada5 (Figure 1). A record from southern Quebec in the summer of 2017 also exists.69,70 Although these records are believed to represent transient incursions,5 Ae. aegypti is thought to have managed to persist through the winter in other cooler locales as larvae in warm subterranean microenvironments.66
Female Aedes aegypti, photo by Adam Blake. This figure appears in color at www.ajtmh.org.
Citation: The American Journal of Tropical Medicine and Hygiene 107, 2; 10.4269/ajtmh.21-0167
Life history.
Historically, a sylvatic tree-hole breeder that fed on animals, almost all populations of Ae. aegypti now preferentially blood-feed on humans,63,71–73 are adapted to human-altered habitats, and breed in a wide variety of artificial containers such as tires, gutters, vases, and buckets74 as well as indoor and underground aquatic habitats.75 Eggs are deposited near the water surface, hatch when the water level rises, and are resistant to desiccation.72,76,77 They are aggressive biters that can feed on multiple hosts during a single gonotrophic cycle, increasing the risk of pathogen transmission.78 Adult female Ae. aegypti are primarily diurnal79,80 and readily enter human habitations to seek a blood meal or rest72; however, they are weak fliers and don’t often fly more than a few hundred meters from breeding sites unless inadvertently transported by humans.81
Taxonomy and identification.
Aedes aegypti is a small black mosquito with stripes of white scales on the tarsomeres and a lyre-shaped pattern of white scales on the scutum74,82 (Figure 2). It looks similar to Ae. sierrensis, Ae. albopictus, Ae. japonicus, and Orthopodomyia spp.; however, the presence of white scales on both the base and apex of tarsomeres of Ae. sierrensis, prominent longitudinal middorsal white stripe of Ae. albopictus, the bronze-scaled lyre-shaped pattern on the scutum of Ae. japonicus, and the lack of distinct stripes of white scales on the fore tarsi of Orthopodomyia spp. can be used to separate these species from Ae. aegypti.
Collection records of Aedes aegypti in Canada.5,70 Note that these records are of ephemeral introductions that are not believed to represent established populations. This figure appears in color at www.ajtmh.org.
Citation: The American Journal of Tropical Medicine and Hygiene 107, 2; 10.4269/ajtmh.21-0167
Public health and veterinary importance.
Aedes aegypti is thus far only believed to present in Canada on a seasonal basis during the summer months.5 However, Ae. aegypti is a principal vector of several important arboviruses including DENV,72 YFV,72 ZIKV,83 and CHIKV.84 The CHIKV and ZIKV are among the most frequent travel-acquired pathogens in Canada,9,85,86 and the presence of Ae. aegypti in Canada, even on an ephemeral basis, raises concerns for autochthonous seasonal transmission of imported arboviruses.
Aedes albopictus distribution.
The Asian tiger mosquito, Aedes (Stegomyia) albopictus (Skuse), is native to southeast Asia but has expanded is range to include an almost global occupation of tropical to temperate habitat.67 Its intercontinental dispersal is thought to be largely due to its use of used tires as breeding habitat,87,88 and within continents it is likely to spread via human-assisted means such as car travel.89
Aedes albopictus was first detected breeding in the United States in 198590 and it has now spread throughout much of the continental United States.67,68 This species was documented in Southern Ontario in 20022 and was found to be established in 2019.5 Aedes albopictus is currently only known in Canada from extreme Southern Ontario (Figure 3), but it has been intercepted in used tires in Seattle,91 not far from the British Columbia border and climate models predict it could establish in several other parts of southern Canada including Quebec, the Prairies, and the southern Maritimes.68 Due to the propensity of Ae. albopictus to spread via human-assisted dispersal and the concentration of the Canadian population in areas that contain suitable climate, or climate that is predicted to become suitable, for Ae. albopictus, this species may spread further within Canada. However, this spread may be slow, sporadic, or limited due to the climatic conditions that are currently marginal for its survival.
Female Aedes albopictus, photo by Ary Faraji. This figure appears in color at www.ajtmh.org.
Citation: The American Journal of Tropical Medicine and Hygiene 107, 2; 10.4269/ajtmh.21-0167
Life history.
Aedes albopictus is an opportunistic daytime blood-feeder on a wide range of animals74,92 that is found in a variety of environments. Aedes albopictus is very flexible in its use of breeding habitat, using a wide variety of artificial containers and even natural habitats like tree holes.74 The larvae of Ae. albopictus are often able to outcompete the larvae of other species that share breeding habitats with them.93,94 At the northern limit of its range in the United States, Ae. albopictus is more abundant in areas with higher mean winter temperatures and March precipitation.95 Diapausing Ae. albopictus eggs that are cold-acclimated can survive at −10°C for up to 24 hours,96 surviving winters that reach lows of −10°C with apparent ease.97 How Ae. albopictus overwinters at the northern limits of its range are unknown, although it has been speculated that this mosquito may have adapted its life history strategy to overwinter as adults in human-made artificial habitats that provide warm microhabitat.5 The survival of eggs or larvae in similar human-made habitats is another possibility, as is a physiological adaptation. Research in this area would be useful in understanding the invasion ecology of this important species.
Taxonomy and identification.
Aedes albopictus is a dark mosquito with bands of white scales on its tarsomeres, an entirely white-scaled 5th hind tarsomere, and a distinct longitudinal stripe of white middorsal scales on its scutum74 (Figure 4). Aedes japonicus and Ae. aegypti can be mistaken for Ae. albopictus due to similar patterns of silvery-white and black scales on the legs, but these species have patterns of scales on the scutum that differ from the middorsal white stripe of Ae. albopictus.
Collection records of Aedes albopictus in Canada.5 Only the population in Windsor, ON, is reported to be established. Other records are of ephemeral introductions that are not believed to represent established populations. This figure appears in color at www.ajtmh.org.
Citation: The American Journal of Tropical Medicine and Hygiene 107, 2; 10.4269/ajtmh.21-0167
Public health and veterinary importance.
Aedes albopictus is a very serious public health concern in much of the world due to its vector competency for exotic arboviruses like ZIKV,98 CHIKV,99 DENV,100 and YFV,101,102 as well as endemic arboviruses like WNV,22 EEEV,103,104 or emerging North American arboviral threats that may spread from the United States to Canada, such as LACV.9 Chikungunya and ZIKV are the common travel-acquired pathogens in Canada,85,86,105 and the establishment of Ae. albopictus may allow for imported cases to result in short-lived autochthonous arbovirus outbreaks, a pattern that occurs in other temperate locations106–108 and has occurred in Canada with other pathogens.43 Aedes albopictus is also a potential vector of dog heartworm.19
Aedes japonicus distribution.
The Asian bush mosquito, Aedes (Hulecoeteomyia) japonicus (Theobald), is endemic to southeast Asia,109 but has expanded its global range drastically over the last several decades. There are four subspecies of Ae. japonicus (Ae. j. japonicus, Ae. j. shintiensis, Ae. j. yaeyamensis, and Ae. j. amamiensis)109 of which Ae. j. japonicus is the only one currently known from North America.110 Since its introduction to North America in 1998111 Ae. j. japonicus has spread over much of the continent and is projected to continue spreading.112 In Canada, it is now known from British Columbia113 including Vancouver Island,114 Ontario,115,116 Quebec,117,118 Newfoundland,119 New Brunswick,120 and Nova Scotia (J. Ogden, pers. comm) (Figure 5).
Female Aedes japonicus, photo by Sean McCann. This figure appears in color at www.ajtmh.org.
Citation: The American Journal of Tropical Medicine and Hygiene 107, 2; 10.4269/ajtmh.21-0167
Genetic evidence indicates that Ae. j. japonicus arrived in eastern North America via at least two separate introductions from Europe,121 and similar studies suggest that populations in western North America are derived from these populations.122 Evidence from the United States suggests that human-assisted dispersal is the main mode of Ae. j. japonicus expansion within North America.123
Suitable habitat exists for Ae. j. japonicus on Prince Edward Island112 and it may only be a matter of time until this species arrives there. Climate change may also lead to suitable habitat for Ae. japonicus in Western Ontario and parts of the Prairies over the coming decades.112 Due to its propensity to spread,124 its mode of spread, the existence of unoccupied suitable habitat, and projections for an increase in suitable habitat over the coming decades, Ae. j. japonicus is likely to continue to spread within Canada.
Life history.
Aedes j. japonicus breeds in tree holes, rock pools, and human containers such as used tires, bird baths, and discarded buckets,110 often containing decaying organic matter, and its distribution is correlated with forested or bushy areas.125 Adults are diurnal and crepuscular in their biting activity and take blood from mammals, including humans, and also from birds.110 Aedes japonicus is tolerant of cold temperatures, emerging earlier and active later in the season than other species that occupy similar ecological niches.109,126 Aedes j. japonicus can overwinter in the egg stage or the larval stage depending on climate and can produce multiple generations per year.110 Warm summer temperatures may prevent Ae. j. japonicus colonization in some areas.127 Laboratory studies have shown that eggs cannot survive extended exposure to temperatures lower than −9°C and that larvae cannot develop at temperatures greater than 31°C.
While there is evidence for competition between Ae. j. japonicus and other mosquitoes altering the assemblage of mosquito species in some areas,127 results from southern Ontario found no change in the species assemblage of mosquitoes with the arrival of Ae. j. japonicus,116 perhaps indicating such effects only occur under specific ecological contexts or at certain scales.
Taxonomy and identification.
A dark mosquito with bands of pale scales on the tarsomeres and a lyre-shaped pattern of bronze scales on the scutum (Figure 6). Adults of Ae. j. japonicus can be distinguished from other Canadian species by the lack of a basal band of pale scales on hind tarsomere 4, dark-scaled abdominal tergites, and a pedicel that usually has more pale scales than dark scales.2,109 A closely related species, Ae. koreicus, has established in Europe,128 outside of its native range in Asia, and maybe an invasive species of concern to Canada. Aedes togoi can be mistaken for Ae. japonicus due to similar patterns of scales on the legs and scutum, but the bands of pale scales on the tarsomeres of Ae. japonicus are present only at the base of each tarsomere.
Collection records of Aedes japonicus in Canada.114–120,219,220 This species is established and widespread in southern Canada outside of drier areas. This figure appears in color at www.ajtmh.org.
Citation: The American Journal of Tropical Medicine and Hygiene 107, 2; 10.4269/ajtmh.21-0167
Public health and veterinary importance.
Aedes japonicus is an aggressive biter that has been implicated as a vector of JEV in its native range,54 and may be a competent vector of WNV,22 EEEV,104 SLEV,129 RVFV,130 DENV and CHIKV,131 and CVV.132 This species is believed to be a significant vector of LACV in parts of the United States,133,134 and it has a high WNV transmission potential in the laboratory135 though field-caught specimens in Ontario rarely test positive for WNV.24
Life history.
Aedes togoi breeds in coastal rock pools just above the high tide line that range from freshwater to hypersaline.140 In Asia, it has also been reported to breed in artificial containers109 but this behavior is absent in North America.140,141 Aedes togoi overwinters in the larval stage, the egg stage, or a combination of both depending on the climate,142,143 with populations in North Vancouver overwintering as a combination of both.144 North American populations do not generally fly more than 20 m from the shoreline.141 Female Ae. togoi will blood-feed on humans,109 but in North America they are often found in locations that are difficult to access and have no human settlements nearby, indicating that their primary source of blood meals are likely other species.140 Due to the reluctance of Ae. togoi adults to leave shoreline habitat,141 a propensity of Ae. togoi larvae to submerge at the slightest provocation and remain hidden in detritus at the bottom of pools for extended periods of time,140 and the nature of its habitat leading to difficulties in access, the detection of Ae. togoi can prove difficult.
Distribution.
Aedes togoi is distributed along the coast of east Asia from Malaysia to the Russian Far East in environments that range from subtropics to subarctic.109 It is also found in Pacific Canada along the coast of southern British Columbia (Figure 7), and down into Washington, though the northern extent of its range in North America is unknown.112,137,138 Aedes togoi was first detected in North America from Victoria, British Columbia, in 1970,139,145 though atypical records from coastal rock pools as early as 1919 may indicate Ae. togoi was observed much earlier but mistaken for other species.138,146 Genetic evidence indicates that populations of Ae. togoi in British Columbia belong to a haplotype not found in known populations from Japan, China, or southeast Asia,136 implying that they may have originated from populations in Primorsky Krai, Russia, or unknown populations elsewhere in the North Pacific.
Female Aedes togoi, photo by Dan Peach. This figure appears in color at www.ajtmh.org.
Citation: The American Journal of Tropical Medicine and Hygiene 107, 2; 10.4269/ajtmh.21-0167
Taxonomy and identification.
Another dark mosquito with banded legs, Ae. togoi has lines of gold scales on the scutum (Figure 8) and is one of the few mosquitoes commonly found in its extreme coastal habitat. Aedes japonicus can be mistaken for Ae. togoi due to similar patterns of scales on the legs and scutum, but the bands of pale scales on the tarsomeres of Ae. togoi are present at the base and apex of each tarsomere, whereas those of Ae. japonicus are present only at the base of each tarsomere.2,82
Collection records of Aedes togoi in Canada.112,139 This species is distributed along the south coast of British Columbia. The lack of records of Ae. togoi from the north coast of British Columbia may represent an information deficiency rather than true absence. This figure appears in color at www.ajtmh.org.
Citation: The American Journal of Tropical Medicine and Hygiene 107, 2; 10.4269/ajtmh.21-0167
Public health and veterinary importance.
In Asia, Ae. togoi is a vector of the filarial parasite Brugia malayi,109,147,148 JEV,53,149,150 and potentially the parasites Wuchereria bancrofti and Dirofilaria immitis.109 However, except in some specific locations, it seems to be secondary in importance to other vector species.
Culex pipiens life history.
The northern house mosquito, Culex pipiens, can breed in almost any type of standing water, particularly containers such a birdbaths, gutters, or buckets, as well as nutrient-rich, stagnant, or polluted water such as ditches or even sewage ponds.4,151,152 Eggs are deposited as a floating raft on the water’s surface, and larvae can develop into adults in 1–2 weeks.4,151 Adult Cx. pipiens are somewhat variable in their hours of peak biting activity but are generally crepuscular to nocturnal.153 This species is primarily ornithophilic, though they do occasionally bite mammals, including humans.20
Culex pipiens numbers build up over the summer, reaching their peak in late July and early August before tapering off into the fall.116 They feed on nectar from a variety of flowers154 and can pollinate some members of the Asteraceae.155 Overwintering occurs as sugar-fed, inseminated nonblood-fed females, which enter warm locations such as human buildings or urban storm water drains in the fall to shelter until they emerge in the spring to feed.156,157
Distribution.
Culex pipiens is a widespread invasive species that has spread throughout the Holarctic as well as into Australia, parts of South America, and South Africa.4,151 In Canada, this species is known from Nova Scotia, New Brunswick, Prince Edward Island, Southern Quebec, Southern Ontario, and British Columbia4,82,114,158,159 (Figure 9). It has been reported from prairie provinces but these records are not supported by specimens or subsequent collections.4,160 Present in North America for centuries, Cx. pipiens perhaps arrived in eastern North America as early as the late fifteenth century.161 However, this species is believed to have spread to Newfoundland only in the early twenty-first century159 and is thought to have been introduced into British Columbia in the early twentieth century, as it was not found in 1904162 and in 1926. Culex pipiens was found only at one location in British Columbia.163 Perhaps, as a result of its use of heated human structures for overwintering, this species is established as far north as Prince George, British Columbia,164 making it the invasive mosquito with the most northernly known distribution in North America. This species had also been reported at Sitka and Yakutat, Alaska, as Cx. consobrinus in the summer of 1899,165 though subsequent Alaskan records have not been reported.82
Female Culex pipiens, photo by Adam Blake. This figure appears in color at www.ajtmh.org.
Citation: The American Journal of Tropical Medicine and Hygiene 107, 2; 10.4269/ajtmh.21-0167
Over the coming decades, environmental conditions are predicted to become suitable for Cx. pipiens in parts of Alberta, Saskatchewan, and Manitoba, as well as Western Ontario.160 Due to its ability to use human structures to escape winter conditions, as well as predictions that future climate conditions will be more favorable to this species in many parts of Canada,160 this species is likely to spread further within Canada.
Taxonomy and identification.
A small brown mosquito without distinct markings (Figure 10), adult female Cx. pipiens can be difficult to reliably distinguish from similar species such as Cx. restuans without molecular tools. However, if specimens are in pristine condition, the presence of patches of pale scales on the scutum can distinguish Cx. restuans from Cx. pipiens.2,4,82 The amphibian biting Cx. territans, another species that appears similar, possesses bands of pale scales on the apices of dorsal abdominal segments, whereas these bands are basal in Cx. pipiens.2,4,82,137 A sister species of Cx. pipiens, the southern house mosquito, Cx. quinquefasciatus, must be separated from Cx. pipiens by molecular techniques or examination of male genitalia; however, Cx. quinquefasciatus is not known from Canada.82
Distribution of Culex pipiens in Canada including the recognized distribution limits of Cx. pipiens in Canada as of 200582 (hash marks), with recent additional collection records.158,159,164,219 The record from near Chisasibi, QC, may not represent an established population and requires further investigation. This figure appears in color at www.ajtmh.org.
Citation: The American Journal of Tropical Medicine and Hygiene 107, 2; 10.4269/ajtmh.21-0167
Public health and veterinary importance.
Though not an aggressive biter,166 Cx. pipiens is an important vector of WNV and is considered as the primary bridge vector of WNV between birds and humans in northeastern North America.20,21,23,24 It is also a vector of SLEV,167,168 USUV,169 avian malaria,170–172 and dog heartworm.173 Similar to human malaria, avian malaria can have massive ecological impacts on the organisms it affects. Declines in populations of the house sparrow, Passer domesticus, in many areas of Europe are thought to be due to Plasmodium relictum174 and the introduction of this parasite into the Hawaiian Islands has devastated endemic bird species, with some experiencing disease mortality of 90% and at least one extinction connected to P. relictum.175–178 The ecology of avian malaria transmission and its impacts need more investigation by vector ecologists, particularly in the northern area of its range and along bird-migration routes.
Future considerations.
Invasive species to watch for.
Aedes atropalpus is native to Eastern North America where it was originally limited to breeding in rock pools.4 Since the late 1970s, it has begun using discarded tires as breeding habitat179 and has become invasive in parts of the American Midwest179 as well as Europe.180,181 By breeding in discarded tires is possible that Ae. atropalpus could invade parts of Canada it has not historically inhabited.
Aedes koreicus is closely related to Ae. japonicus and is abundant in urban settings.182 The native range of Ae. koreicus is in eastern Asia, including China, Korea, and eastern Russia,109 but it has become established in Europe where it is believed to have arrived in used tires.128,183 As a container-breeding species well-adapted to temperate climates, human-dominated environments, and a history of spreading to new areas, Ae. koreicus may have the potential to invade and establish in Canada.
Culex tritaeniorhynchus is native to Asia and parts of Africa109 and is the primary vector of JEV virus in many parts of Asia.109 This species has been found on ships as far as several hundred kilometers out to sea184 and, while it primarily breeds in other habitats, its larvae have occasionally been found in containers.185 Culex tritaeniorhynchus could arrive as ship-borne adults or as larvae in containers shipped from its native habitat.
DISCUSSION
With an increasingly interconnected world, the establishment of invasive alien species is on the rise worldwide.186 At least half of the invasive mosquitoes known from Canada have arrived within the last 20 years, and all but one since the middle of the twentieth century. Mosquito and pathogen monitoring and control in Canada must adapt to these challenges by utilizing modern solutions and undertaking context-specific research.
As genomic resources for mosquitoes continue to improve, the ability to identify species via molecular means will become increasingly important in surveillance and control efforts. Widely deployed molecular identification (e.g., through “DNA barcoding”187) is a critical component of vector surveillance, as it can resolve cryptic or sibling species as well as reduce the need for highly trained observers. Furthermore, molecular identification can be performed on partial or damaged samples from multiple life stages,188 obviating the need for pristine specimens for morphological identification. Pathogen surveillance programs have traditionally relied upon morphological identification techniques, which have resulted in the successful detection of several previously unknown native mosquitoes3,189 and novel invasives.5,113 However, DNA barcoding methods were recently used in a province-wide survey of mosquitoes in Quebec118 and we propose that similar methods should be routinely and widely implemented (in tandem with morphological identification techniques) as part of mosquito surveillance efforts across Canada. In addition to species identification, sequencing analysis can help to identify the likely source of invasive mosquitoes,63,64,190,191 which will aid in targeting surveillance and trapping to detect and prevent further incursion (e.g., at ports or airports). Finally, sequencing intact mosquito specimens can identify arbovirus infection status and bloodmeal host identity73,192–198 providing critical information to aid in control efforts. We propose that the generation of high-quality reference genomes should be prioritized for mosquitoes native and invasive to Canada. Contiguous and complete genome assemblies can now be readily constructed from as little as a single mosquito due to improvements in long-read sequencing technologies and library preparation protocols.199–201 The availability of comprehensive and complete genomic resources will help to better understand the biology of these species and to guide the development of molecular assays for determining insecticide resistance status.202–205 Further, coupled with whole-genome sequencing of additional specimens, a complete genome assembly is a foundation identifying the genomic basis of oviposition behavior, blood-meal host preference, and other phenotypes that may influence the ability of specific populations to take hold and adapt to new niches.206 However, it is important to stress that molecular techniques do not obviate the importance of morphological techniques in vector surveillance and at broader scales,207,208 and that future best practices will involve a reciprocal and integrative interaction between these methodologies.209
Broad collaboration and data-sharing are imperative to monitor, prevent, and prepare for the spread of invasive mosquitoes and the pathogens they vector.210–212 Requirements for open data policies combined with a greater resolution of data, such as species-level identifications, for such institutions that monitor for mosquitoes and pathogens, and their contractors, are one mechanism by which to address these needs. Detailed data collection and open data policies regarding species and pathogen presence will also inform modeling efforts and risk assessments, policy decisions, and healthcare initiatives. To facilitate these efforts, and build upon pathogen monitoring activities such as those carried out by provincial CDC, the Public Health Agency of Canada, Public Health Ontario, and others, we propose the formation of a Canada-wide network for sharing information on the distribution of mosquito vectors and the pathogens they spread, similar to the VectorNet (https://vectornet.ecdc.europa.eu/) and VBORNET (www.vbornet.eu) initiatives established by the European Center for Disease Prevention and Control and European Food Safety Authority. Additionally, monitoring for invasive mosquitoes arriving at seaports, airports, and other areas at which high volumes of international traffic arrive, as has been done in other nations213–216 and in some limited parts of Canada such as southern Ontario,5,26 should be considered to enhance Canada’s biosecurity due to the ease at which some invasive mosquitoes spread within a country once established.91,110,217,218
Furthermore, Canada provides a unique opportunity to study invasive mosquitoes and their pathogens under an environment that is changing in both climate and levels of human modification. Many of these mosquitoes and pathogens are at the northern limits of their range in Canada and are likely experiencing immense selection pressures on behavioral and physiological mechanisms to survive cold winters. With changing climatic conditions and increasing global connectivity, there are also unique opportunities to study how invasive mosquitoes and exotic pathogens arrive and spread in new areas, as well as the effect of urbanization and temperature limitations on mosquito and pathogen community composition, mosquito behavior, and pathogen transmission dynamics.
ACKNOWLEDGMENTS
We thank Adam Blake for the photographs of Ae. aegypti and Cx. pipiens, Sean McCann for the photograph of Ae. japonicus, and Ary Faraji for the photograph of Ae. albopictus. We also thank Public Health Ontario, the Royal BC Museum, and the UBC Beaty Biodiversity Museum for sharing mosquito collection records.
REFERENCES
- 1.↑
Gates B , 2014. The Deadliest Animal in the World. Available at: https://www.gatesnotes.com/Health/Most-Lethal-Animal-Mosquito-Week. Accessed January 26, 2020.
- 2.↑
Thielman A , Hunter F , 2007. Photographic key to the adult female mosquitoes (Diptera: Culicidae) of Canada. Can J Arthropod Identif 4: 1–117.
- 3.↑
Jackson M , Howay T , Belton P , 2013. The first record of Culiseta particeps (Diptera: Culicidae) in Canada. Can Entomol 145: 115–116.
- 4.↑
Wood DM , Dang PT , Ellis RA , 1979. The Insects and Arachnids of Canada Part 6: The Mosquitoes of Canada - Diptera: Culicidae. Ottawa, Canada: Research Branch, Agriculture Canada.
- 5.↑
Giordano BV , Gasparotto A , Liang P , Nelder MP , Russell C , Hunter FF , 2020. Discovery of an Aedes (Stegomyia) albopictus population and first records of Aedes (Stegomyia) aegypti in Canada. Med Vet Entomol 34: 10–16.
- 6.↑
Canadian Council on Invasive Species , 2022. Invasive Species. Available at: https://canadainvasives.ca/invasive-species/. Accessed April 15, 2022.
- 7.↑
Tomasello D , Schlagenhauf P , 2013. Chikungunya and dengue autochthonous cases in Europe, 2007–2012. Travel Med Infect Dis 11: 274–284.
- 8.↑
Medlock JM , Hansford KM , Versteirt V , Cull B , Kampen H , Fontenille D , Hendrickx G , Zeller H , Van Bortel W , Schaffner F , 2015. An entomological review of invasive mosquitoes in Europe. Bull Entomol Res 105: 637–663.
- 9.↑
Drebot M , 2015. Emerging mosquito-borne bunyaviruses in Canada. Can Commun Dis Rep 41: 117–123.
- 10.↑
Artsob H , 1990. Arbovirus activity in Canada. Calisher C , ed. Hemorrhagic Fever with Renal Syndrome, Tick- and Mosquito-Borne Viruses. Archives of Virology Supplementum, Vol. 1. Vienna, Austria: Springer, 249–258.
- 11.↑
Walters LL , Tirrell SJ , Shope RE , 1999. Seroepidemiology of California and Bunyamwera serogroup (Bunyaviridae) virus infections in native populations of Alaska. Am J Trop Med Hyg 60: 806–821.
- 13.↑
Lau L , Wudel B , Kadkhoda K , Keynan Y , 2017. Snowshoe hare virus causing meningoencephalitis in a young adult from northern Manitoba, Canada. Open Forum Infect Dis 4: 514–534.
- 14.↑
Heath SE , Artsob H , Bell RJ , Harland RJ , 1989. Equine encephalitis caused by snowshoe hare (California serogroup) virus. Can Vet J 30: 669–671.
- 15.↑
McCall JW , Genchi C , Kramer LH , Guerrero J , Venco L , 2008. Heartworm disease in animals and humans. Adv Parasitol 66: 193–285.
- 16.↑
Diaz JH , Risher WH , 2015. Risk factors for human heartworm infections (dirofilariasis) in the South. J La State Med Soc 167: 79–86.
- 17.↑
Herrin BH , Peregrine AS , Goring J , Beall MJ , Little SE , 2017. Canine infection with Borrelia burgdorferi, Dirofilaria immitis, Anaplasma spp. and Ehrlichia spp. in Canada, 2013–2014. Parasit Vectors 10: 244.
- 18.↑
McGill E , Berke O , Weese JS , Peregrine A , 2019. Heartworm infection in domestic dogs in Canada, 1977–2016: prevalence, time trend, and efficacy of prophylaxis. Can Vet J 60: 605–612.
- 19.↑
Ledesma N , Harrington L , 2011. Mosquito vectors of dog heartworm in the United States: vector status and factors influencing transmission efficiency. Top Companion Anim Med 26: 178–185.
- 20.↑
Hamer GL , Kitron UD , Brawn JD , Loss SR , Ruiz MO , Goldberg TL , Walker ED , 2008. Culex pipiens (Diptera: Culicidae): a bridge vector of West Nile virus to humans. J Med Entomol 45: 125–128.
- 21.↑
Hamer GL , Kitron UD , Goldberg TL , Brawn JD , Loss SR , Ruiz MO , Hayes DB , Walker ED , 2009. Host selection by Culex pipiens mosquitoes and West Nile virus amplification. Am J Trop Med Hyg 80: 268–278.
- 22.↑
Turell MJ , Dohm DJ , Sardelis MR , O’Guinn ML , Andreadis TG , Blow JA , 2005. An update on the potential of North American mosquitoes (Diptera: Culicidae) to transmit West Nile virus. J Med Entomol 42: 6.
- 23.↑
Andreadis TG , 2012. The contribution of Culex pipiens complex mosquitoes to transmission and persistence of West Nile virus in North America. Moco 28: 137–151.
- 24.↑
Giordano BV , Kaur S , Hunter FF , 2017. West Nile virus in Ontario, Canada: a twelve-year analysis of human case prevalence, mosquito surveillance, and climate data. PLoS One 12: e0183568.
- 25.↑
Roth D , Henry B , Mak S , Fraser M , Taylor M , Li M , Cooper K , Furnell A , Wong Q , Morshed M , 2010. West Nile virus range expansion into British Columbia. Emerg Infect Dis 16: 1251–1258.
- 26.↑
Giordano B , Turner K , Hunter F , 2018. Geospatial analysis and seasonal distribution of West Nile virus vectors (Diptera: Culicidae) in southern Ontario, Canada. IJERPH 15: 614.
- 27.↑
Kilpatrick AM , Kramer LD , Jones MJ , Marra PP , Daszak P , 2006. West Nile virus epidemics in North America are driven by shifts in mosquito feeding behavior. PLoS Biol 4: e82.
- 28.↑
Nash D et al.2001. The outbreak of West Nile virus infection in the New York City area in 1999. N Engl J Med 344: 1807–1814.
- 29.↑
Drebot MA , Lindsay R , Barker IK , Buck PA , Fearon M , Hunter F , Sockett P , Artsob H , 2003. West Nile virus surveillance and diagnostic: a Canadian perspective. Can J Infect Dis 14: 105–114.
- 31.↑
Eidson M , Komar N , Sorhage F , Nelson R , Talbot T , Mostashari F , McLean R , 2001. Crow deaths as a sentinel surveillance system for West Nile virus in the northeastern United States, 1999. Emerg Infect Dis 7: 6.
- 32.↑
Calisher CH , 1994. Medically important arboviruses of the United States and Canada. Clin Microbiol Rev 7: 28.
- 33.↑
Flores FS , Diaz LA , Batallán GP , Almirón WR , Contigiani MS , 2010. Vertical transmission of St. Louis encephalitis virus in Culex quinquefasciatus (Diptera: Culicidae) in Córdoba, Argentina. Vector Borne Zoonotic Dis 10: 999–1002.
- 34.↑
Rosa R , Costa EA , Marques RE , Oliveira TS , Furtini R , Bomfim MRQ , Teixeira MM , Paixão TA , Santos RL , 2013. Isolation of Saint Louis encephalitis virus from a horse with neurological disease in Brazil. PLoS Negl Trop Dis 7: e2537.
- 35.↑
Andreadis TG , Anderson JF , Armstrong PM , Main AJ , 2008. Isolations of Jamestown Canyon virus (Bunyaviridae: Orthobunyavirus) from field-collected mosquitoes (Diptera: Culicidae) in Connecticut, USA: a ten-year analysis, 1997–2006. Vector Borne Zoonotic Dis 8: 175–188.
- 36.↑
Pastula DM , Hoang Johnson DK , Fischer M , White JL , Staples JE , Dupuis AP , 2015. Jamestown Canyon virus disease in the United States—2000–2013. Am J Trop Med Hyg 93: 384–389.
- 37.↑
Iversen JO , Wagner RJ , Leung MK , Hayles LB , McLintock JR , 1979. Cache Valley virus: isolations from mosquitoes in Saskatchewan, 1972–1974. Can J Microbiol 25: 760–764.
- 38.↑
Nguyen NL , Zhao G , Hull R , Shelly MA , Wong SJ , Wu G , St George K , Wang D , Menegus MA , 2013. Cache valley virus in a patient diagnosed with aseptic meningitis. J Clin Microbiol 51: 1966–1969.
- 39.↑
Chung SI , Livingston CW , Edwards JF , Crandell RW , Shope RE , Shelton MJ , Collisson EW , 1990. Evidence that Cache Valley virus induces congenital malformations in sheep. Vet Microbiol 21: 297–307.
- 40.↑
Harvey J , Smith J , Jackson N , Kreuder A , Dohlman T , Smith J , 2019. Cache Valley virus as a cause of fetal abnormalities in a litter of three Boer kids. Vet Rec Case Rep 7: e000725.
- 41.↑
Ozburn RH , 1944. Preliminary report on anopheline mosquito survey in Canada. Pt. 1. A report on light trap collections. Entomological Society of Ontario Annual Report 75: 37–44.
- 42.↑
Hadwin S , 1915. A note on the occurrence and significance of Anophelinae in B.C. J Entomol Soc BC 5: 81–82.
- 43.↑
Ndao M , Bandyayera E , Kokoskin E , Gyorkos TW , MacLean JD , Ward BJ , 2004. Comparison of blood smear, antigen detection, and nested-PCR methods for screening refugees from regions where malaria is endemic after a malaria outbreak in Quebec, Canada. J Clin Microbiol 42: 2694–2700.
- 44.↑
Rocheleau J-P , Arsenault J , Lindsay LR , Dibernardo A , Kulkarni MA , Côté N , Michel P , 2013. Eastern equine encephalitis virus: high seroprevalence in horses from southern Quebec, Canada, 2012. Vector Borne Zoonotic Dis 13: 712–718.
- 45.↑
Molaei G , Oliver J , Andreadis TG , Armstrong PM , Howard JJ , 2006. Molecular identification of blood-meal sources in Culiseta melanura and Culiseta morsitans from an endemic focus of eastern equine encephalitis virus in New York. Am J Trop Med Hyg 75: 1140–1147.
- 46.↑
Nelder MP , Moore S , Russell C , Sider D , Ontario Agency for Health Protection and Promotion , 2014. Eastern Equine Encephalitis Virus: History and Enhanced Surveillance in Ontario. Toronto, ON: Queen’s Printer for Ontario.
- 47.↑
Wages DP , Ficken MD , Guy JS , Cummings TS , Jennings SR , 1993. Egg-production drop in turkeys associated with alphaviruses: eastern equine encephalitis virus and Highlands J virus. Avian Dis 37: 1163.
- 48.↑
Cooper GL , Medina HA , 1999. Egg production drops in breeder turkeys associated with western equine encephalitis virus infection. Avian Dis 43: 136.
- 50.↑
Marx PA , Hanley KA , Weaver SC , Vasilakis N , Whitehead SS , Brown M , Kautz TF , Guerbois M , 2014. Infection dynamics of sylvatic dengue virus in a natural primate host, the African green monkey. Am J Trop Med Hyg 91: 672–676.
- 51.↑
Screaton G , Mongkolsapaya J , Yacoub S , Roberts C , 2015. New insights into the immunopathology and control of dengue virus infection. Nat Rev Immunol 15: 745–759.
- 52.↑
van den Hurk AF , Ritchie SA , Mackenzie JS , 2009. Ecology and geographical expansion of Japanese encephalitis virus. Annu Rev Entomol 54: 17–35.
- 53.↑
Petrishcheva P , 1948. On the transition of Aedes togoi Theob. and Aedes japonicus Theob. (Diptera, Culicidae) to a synanthropic form of life. Entomol Obozr 30: 103–108.
- 54.↑
Takashima I , Rosen L , 1989. Horizontal and vertical transmission of Japanese encephalitis virus by Aedes japonicus (Diptera: Culicidae). J Med Entomol 26: 454–458.
- 55.↑
Nikolay B , Diallo M , Boye CSB , Sall AA , 2011. Usutu virus in Africa. Vector Borne Zoonotic Dis 11: 1417–1423.
- 56.↑
Pecorari M et al.2009. First human case of Usutu virus neuroinvasive infection, Italy, August–September 2009. Euro Surveill 14: 19446.
- 57.↑
Douam F , Ploss A , 2018. Yellow fever virus: knowledge gaps impeding the fight against an old foe. Trends Microbiol 26: 913–928.
- 58.↑
Dorigatti I , Hamlet A , Aguas R , Cattarino L , Cori A , Donnelly CA , Garske T , Imai N , Ferguson NM , 2017. International risk of yellow fever spread from the ongoing outbreak in Brazil, December 2016 to May 2017. Euro Surveill 22: 30572.
- 60.↑
Silva JVJ , Ludwig-Begall LF , de Oliveira-Filho EF , Oliveira RAS , Durães-Carvalho R , Lopes TRR , Silva DEA , Gil LHVG , 2018. A scoping review of Chikungunya virus infection: epidemiology, clinical characteristics, viral co-circulation complications, and control. Acta Trop 188: 213–224.
- 61.↑
Chompoosri J , Thavara U , Tawatsin A , Boonserm R , Phumee A , Sangkitporn S , Siriyasatien P , 2016. Vertical transmission of Indian Ocean Lineage of chikungunya virus in Aedes aegypti and Aedes albopictus mosquitoes. Parasit Vectors 9: 227.
- 62.↑
Fawzy M , Helmy YA , 2019. The One Health approach is necessary for the control of Rift Valley fever infections in Egypt: a comprehensive review. Viruses 11: 139.
- 63.↑
Powell JR , Tabachnick WJ , 2013. History of domestication and spread of Aedes aegypti—a review. Mem Inst Oswaldo Cruz 108: 11–17.
- 64.↑
Powell JR , Gloria-Soria A , Kotsakiozi P , 2018. Recent history of Aedes aegypti: vector genomics and epidemiology records. Bioscience 68: 854–860.
- 65.↑
Jansen CC , Beebe NW , 2010. The dengue vector Aedes aegypti: what comes next. Microbes Infect 12: 272–279.
- 66.↑
Lima A , Lovin DD , Hickner PV , Severson DW , 2016. Evidence for an overwintering population of Aedes aegypti in Capitol Hill neighborhood, Washington, DC. Am J Trop Med Hyg 94: 231–235.
- 67.↑
Kraemer MUG et al.2015. The global distribution of the arbovirus vectors Aedes aegypti and Ae. albopictus. eLife 4: 1–18.
- 68.↑
Kraemer MUG et al.2019. Past and future spread of the arbovirus vectors Aedes aegypti and Aedes albopictus. Nat Microbiol 4: 854–863.
- 69.↑
Ludwig A , Zheng H , Vrbova L , Drebot M , Iranpour M , Lindsay L , 2019. Increased risk of endemic mosquito-borne diseases in Canada due to climate change. CCDR 45: 91–97.
- 70.↑
Lapierre M. , 2018. Chance of contracting the Zika virus in Quebec “is negligible to nul.” June 13, 2018. Montreal Gazette. Available at: https://montrealgazette.com/news/quebec/chance-of-contracting-the-zika-virus-in-quebec-is-negligible-to-nul. Accessed May 19, 2020.
- 71.↑
McBride CS , Baier F , Omondi AB , Spitzer SA , Lutomiah J , Sang R , Ignell R , Vosshall LB , 2014. Evolution of mosquito preference for humans linked to an odorant receptor. Nature 515: 222–227.
- 72.↑
Christophers SR , 1960. Aedes aegypti (L.) the Yellow Fever Mosquito: Its Life History, Bionomics and Structure. Cambridge, United Kingdom: Cambridge University Press.
- 73.↑
Ponlawat A , Harrington LC , 2005. Blood feeding patterns of Aedes aegypti and Aedes albopictus in Thailand. J Med Entomol 42: 844–849.
- 74.↑
Burkett-Cadena ND , 2013. Mosquitoes of the Southeastern United States. Tuscaloosa, AL: University of Alabama Press.
- 75.↑
Barrera R , Amador M , Diaz A , Smith J , Munoz-Jordan JL , Rosario Y , 2008. Unusual productivity of Aedes aegypti in septic tanks and its implications for dengue control. Med Vet Entomol 22: 62–69.
- 76.↑
Juliano SA , Philip Lounibos L , 2005. Ecology of invasive mosquitoes: effects on resident species and on human health: invasive mosquitoes. Ecol Lett 8: 558–574.
- 77.↑
Carpenter S , LaCasse W , 1955. Mosquitoes of North America (North of Mexico). Berkeley, CA: University of California Press.
- 78.↑
Farjana T , Tuno N , 2013. Multiple blood feeding and host-seeking behavior in Aedes aegypti and Aedes albopictus (Diptera: Culicidae). J Med Entomol 50: 838–846.
- 79.↑
Araujo M da S , Guo F , Rosbash M , 2020. Video recording can conveniently assay mosquito locomotor activity. Sci Rep 10: 4994.
- 80.↑
Yee WL , Foster WA , 1992. Diel sugar-feeding and host-seeking rhythms in mosquitoes (Diptera: Culicidae) under laboratory conditions. J Med Entomol 29: 784–791.
- 81.↑
Verdonschot PFM , Besse-Lototskaya AA , 2014. Flight distance of mosquitoes (Culicidae): a metadata analysis to support the management of barrier zones around rewetted and newly constructed wetlands. Limnologica 45: 69–79.
- 82.↑
Darsie RFJ , Ward RA , 2005. Identification and Geographical Distribution of the Mosquitoes of North America, North of Mexico. Gainesville, FL: University Press of Florida.
- 83.↑
Ferreira-de-Brito A , Ribeiro IP , de Miranda RM , Fernandes RS , Campos SS , da Silva KAB , de Castro MG , Bonaldo MC , Brasil P , Lourenço-de-Oliveira R , 2016. First detection of natural infection of Aedes aegypti with Zika virus in Brazil and throughout South America. Mem Inst Oswaldo Cruz 111: 655–658.
- 84.↑
Vega-Rúa A , Zouache K , Girod R , Failloux A-B , Lourenço-de-Oliveira R , 2014. High level of vector competence of Aedes aegypti and Aedes albopictus from ten American countries as a crucial factor in the spread of chikungunya virus. J Virol 88: 6294–6306.
- 85.↑
Tataryn J , Vrbova L , Drebot M , Wood H , Payne E , Connors S , Geduld J , German M , Khan K , Buck P , 2018. Travel-related Zika virus cases in Canada: October 2015–June 2017. Can Commun Dis Rep 44: 18–26.
- 86.↑
Ng V , Rees E , Lindsay R , Drebot M , Brownstone T , Sadeghieh T , Khan S , 2019. Could exotic mosquito-borne diseases emerge in Canada with climate change? CCDR 45: 98–107.
- 87.↑
Hawley W , Reiter P , Copeland R , Pumpuni C , Craig G , 1987. Aedes albopictus in North America: probable introduction in used tires from northern Asia. Science 236: 1114–1116.
- 88.↑
Bennett KL et al.2019. High infestation of invasive Aedes mosquitoes in used tires along the local transport network of Panama. Parasit Vectors 12: 264.
- 89.↑
Eritja R , Palmer JRB , Roiz D , Sanpera-Calbet I , Bartumeus F , 2017. Direct evidence of adult Aedes albopictus dispersal by car. Sci Rep 7: 14399.
- 90.↑
Sprenger D , Wuithiranyagool T , 1986. The discovery and distribution of Aedes albopictus in Harris County, Texas. J Am Mosq Control Assoc 2: 217–219.
- 91.↑
Moore CG , 1999. Aedes albopictus in the United States: current status and prospects for futher spread. J Am Mosq Control Assoc 15: 221–227.
- 92.↑
Niebylski ML , Savage HM , Nasci RS , Craig GB Jr , 1994. Blood hosts of Aedes albopictus in the United States. J Am Mosq Control Assoc 10: 447–450.
- 93.↑
Livdahl TP , Willey MS , 1991. Prospects for an invasion: competition between Aedes albopictus and native Aedes triseriatus. Science 253: 189–191.
- 94.↑
Costanzo KS , Mormann K , Juliano SA , 2005. Asymmetrical competition and patterns of abundance of Aedes albopictus and Culex pipiens (Diptera: Culicidae). J Med Entomol 42: 12.
- 95.↑
Kache PA , Eastwood G , Collins-Palmer K , Katz M , Falco RC , Bajwa WI , Armstrong PM , Andreadis TG , Diuk-Wasser MA , 2020. Environmental determinants of Aedes albopictus abundance at a northern limit of its range in the United States. Am J Trop Med Hyg 102: 436–447.
- 96.↑
Thomas SM , Obermayr U , Fischer D , Kreyling J , Beierkuhnlein C , 2012. Low-temperature threshold for egg survival of a post-diapause and non-diapause European aedine strain, Aedes albopictus (Diptera: Culicidae). Parasit Vectors 5: 100.
- 97.↑
Tippelt L , Werner D , Kampen H , 2019. Tolerance of three Aedes albopictus strains (Diptera: Culicidae) from different geographical origins towards winter temperatures under field conditions in northern Germany. PLoS One 14: e0219553.
- 98.↑
Grard G , Caron M , Mombo IM , Nkoghe D , Mboui Ondo S , Jiolle D , Fontenille D , Paupy C , Leroy EM , 2014. Zika virus in Gabon (Central Africa)—2007: a new threat from Aedes albopictus? PLoS Negl Trop Dis 8: e2681.
- 99.↑
Zouache K , Michelland RJ , Failloux A-B , Grundmann GL , Mavingui P , 2012. Chikungunya virus impacts the diversity of symbiotic bacteria in mosquito vector. Mol Ecol 21: 2297–2309.
- 100.↑
Delatte H , Paupy C , Dehecq JS , Thiria J , Failloux AB , Fontenille D , 2008. Aedes albopictus, vecteur des virus du chikungunya et de la dengue à la Réunion: biologie et contrôle. Parasite 15: 3–13.
- 101.↑
Mitchell CJ , Miller BR , Gubler DJ , 1987. Vector competence of Aedes albopictus from Houston, Texas, for dengue serotypes 1 to 4, yellow fever and Ross River viruses. J Am Mosq Control Assoc 3: 460–465.
- 102.↑
Amraoui F , Vazeille M , Failloux AB , 2016. French Aedes albopictus are able to transmit yellow fever virus. Euro Surveill 21: 30361.
- 103.↑
Turell MJ , Beaman JR , Neely GW , 1994. Experimental transmission of eastern equine encephalitis virus by strains of Aedes albopictus and A. taeniorhynchus (Diptera: Culicide). J Med Entomol 31: 287–290.
- 104.↑
Sardelis MR , Dohm DJ , Pagac B , Andre RG , Turell MJ , 2002. Experimental transmission of eastern equine encephalitis virus by Ochlerotatus j. japonicus (Diptera: Culicidae). J Med Entomol 39: 480–484.
- 105.↑
Drebot M , Holloway K , Zheng H , Ogden N , 2015. Travel-related chikungunya cases in Canada, 2014. CCDR 41: 2–5.
- 106.↑
Rezza G et al.2007. Infection with chikungunya virus in Italy: an outbreak in a temperate region. Lancet 370: 1840–1846.
- 107.↑
Kutsuna S et al.2015. Autochthonous dengue fever, Tokyo, Japan, 2014. Emerg Infect Dis 21: 517–520.
- 108.↑
Septfons A et al.2016. Travel-associated and autochthonous Zika virus infection in mainland France, 1 January to 15 July 2016. Euro Surveill 21: 30315.
- 109.↑
Tanaka K , Mizusawa K , Saugstad ES , 1979. A revision of the adult and larval mosquitoes of Japan (including the Ryukyu Archipelago and the Ogasawara Islands) and Korea (Diptera: Culicidae). Contrib Am Entomol Inst 16: 989.
- 110.↑
Kaufman MG , Fonseca DM , 2014. Invasion biology of Aedes japonicus japonicus (Diptera: Culicidae). Annu Rev Entomol 59: 31–49.
- 111.↑
Peyton EL , Campbell SR , Candeletti TM , Romanoski M , Crans WJ , 1999. Aedes (Finlaya) japonicus japonicus (Theobald), a new introduction into the United States. J Am Mosq Control Assoc 15: 238–241.
- 112.↑
Peach DAH , Almond M , Pol J , 2019. Modeled distributions of Aedes japonicus and Aedes togoi (Diptera: Culicidae) in the United States, Canada, and northern Latin America. J Vector Ecol 44: 119–129.
- 113.↑
Jackson M , Belton P , McMahon S , Hart M , McCann S , Azevedo D , Hurteau L , 2016. The first record of Aedes (Hulecoeteomyia) japonicus (Diptera: Culicidae) and its establishment in Western Canada. J Med Entomol 53: 241–244.
- 114.↑
Peach DAH , 2018. An updated list of the mosquitoes of British Columbia with distribution notes. J Entomol Soc BC 115: 126–129.
- 115.↑
Thielman A , Hunter F , 2006. Establishment of Ochlerotatus japonicus (Diptera: Culicidae) in Ontario, Canada. J Med Entomol 43: 138–142.
- 116.↑
Dussault C , Nelder MP , Russell C , Johnson S , Vrbova L , 2018. Evaluating the impact of Aedes japonicus invasion on the mosquito community in the Greater Golden Horseshoe region (Ontario, Canada). PLoS One 13: e0208911.
- 117.↑
Savignac R , Back C , Bourassa J , 2002. Biological notes on Ochlerotatus japonicus and other mosquito species new to Quebec. Joint Meeting 68th Annual Meeting of the American Mosquito Control Association and the West Central Mosquito and Vector Control Association, 21–22, Denver, CO.
- 118.↑
Shahhosseini N , Wong G , Frederick C , Kobinger GP , 2020. Mosquito species composition and abundance in Quebec, eastern Canada. J Med Entomol 57: tjaa020.
- 119.↑
Fielden MA , Chaulk AC , Bassett K , Wiersma YF , Erbland M , Whitney H , Chapman TW , 2015. Aedes japonicus japonicus (Diptera: Culicidae) arrives at the most easterly point in North America. Can Entomol 147: 737–740.
- 120.↑
Edsall J , Webster RP , Giguère M-A , Maltais P , Roy J , 2010. Mosquitoes (Diptera: Culicidae) of the Atlantic Maritime ecozone. McAlpine DF , Smith IM , eds. Assessment of Species Diversity in the Atlantic Maritime Ecozone. Ottawa, ON: NRC Research Press.
- 121.↑
Fonseca DM , Widdel AK , Hutchinson M , Spichiger SE , Kramer LD , 2010. Fine-scale spatial and temporal population genetics of Aedes japonicus, a new US mosquito, reveal multiple introductions. Mol Ecol 19: 1559–1572.
- 122.↑
Baharmand I , Coatsworth H , Peach DAH , Belton P , Lowenberger C , 2020. Exploring the molecular relationships and potential origins of introduced Aedes japonicus (Diptera: Culicidae) populations in British Columbia, Canada using mitochondrial DNA. J Vector Ecol 45: 285–296.
- 123.↑
Egizi A , Kiser J , Abadam C , Fonseca DM , 2016. The hitchhiker’s guide to becoming invasive: exotic mosquitoes spread across a US state by human transport not autonomous flight. Mol Ecol 25: 3033–3047.
- 124.↑
Montarsi F et al.2019. The invasive mosquito Aedes japonicus japonicus is spreading in northeastern Italy. Parasit Vectors 12: 120.
- 125.↑
Bartlett-Healy K et al.2012. Larval mosquito habitat utilization and community dynamics of Aedes albopictus and Aedes japonicus (Diptera: Culicidae). J Med Entomol 49: 813–824.
- 126.↑
Iriarte WLZ , Tsuda Y , Wada Y , Takagi M , 1991. Distribution of mosquitoes on a hill of Nagasaki City, with emphasis to the distance from human dwellings. Trop Med 33: 55–60.
- 127.↑
Andreadis TG , Wolfe RJ , 2010. Evidence for reduction of native mosquitoes with increased expansion of invasive Ochlerotatus japonicus japonicus (Diptera: Culicidae) in the northeastern United States. J Med Entomol 47: 43–52.
- 128.↑
Versteirt V , Pecor JE , Fonseca DM , Coosemans M , Van Bortel W , 2012. Confirmation of Aedes koreicus (Diptera: Culicidae) in Belgium and description of morphological differences between Korean and Belgian specimens validated by molecular identification. Zootaxa 3191: 21.
- 129.↑
Sardelis MR , Turell MJ , Andre RG , 2003. Experimental transmission of St. Louis encephalitis virus by Ocherotatus j. japonicus. J Am Mosq Control Assoc 19: 159–162.
- 130.↑
Turell MJ , Byrd BD , Harrison BA , 2013. Potential for populations of Aedes j. japonicus to transmit Rift Valley fever virus in the USA. J Am Mosq Control Assoc 29: 133–137.
- 131.↑
Schaffner F , Vazeille M , Kaufmann C , Failloux A , Mathis A , 2011. Vector competence of Aedes japonicus for chikungunya and dengue viruses. Eur Mosq Bull 29: 141–142.
- 132.↑
Yang F , Chan K , Marek PE , Armstrong PM , Liu P , Bova JE , Bernick JN , McMillan BE , Weidlich BG , Paulson SL , 2018. Cache Valley virus in Aedes japonicus japonicus mosquitoes, Appalachian region, United States. Emerg Infect Dis 24: 553–557.
- 133.↑
Harris MC , Dotseth EJ , Jackson BT , Zink SD , Marek PE , Kramer LD , Paulson SL , Hawley DM , 2015. La Crosse virus in Aedes japonicus japonicus mosquitoes in the Appalachian Region, United States. Emerg Infect Dis 21: 646–649.
- 134.↑
Bara JJ , Parker AT , Muturi EJ , 2016. Comparative Susceptibility of Ochlerotatus japonicus, Ochlerotatus triseriatus, Aedes albopictus, and Aedes aegypti (Diptera: Culicidae) to La Crosse virus. J Med Entomol 53: 1415–1421.
- 135.↑
Turell MJ , O’Guinn ML , Dohm DJ , Jones JW , 2001. Vector competence of North American mosquitoes (Diptera: Culicidae) for West Nile virus. J Med Entomol 38: 130–134.
- 136.↑
Sota T , Belton P , Tseng M , Yong HS , Mogi M , 2015. Phylogeography of the coastal mosquito Aedes togoi across climatic zones: testing an anthropogenic dispersal hypothesis. PLoS One 10: e0131230.
- 137.↑
Belton P , 1983. The Mosquitoes of British Columbia. Victoria, BC, Canada: British Columbia Provincial Museum.
- 138.↑
Peach DAH , Matthews BJ , 2020. Modelling the putative ancient distribution of the coastal rock pool mosquito Aedes togoi. J Insect Sci 20: 1–10.
- 139.↑
Belton P , 1980. The first record of Aedes togoi (Theo.) in the United States - aboriginal or ferry passenger? Mosq News 40: 624–626.
- 140.↑
Sames J , Wlliam I , Florin’ A , Francis E , Maloneyi A , 2004. Distribution of Ochlerotatus togoi along the Pacific Coast of Washington. J Am Mosq Control Assoc 20: 105–109.
- 141.↑
Trimble RM , 1984. Aedes togoi (Diptera: Culicidae) dispersal: assessment using artificial container habitats and miniature light traps. J Med Entomol 21: 120–121.
- 142.↑
Sota T , 1994. Larval diapause, size, and autogeny in the mosquito Aedes togoi (Diptera, Culicidae) from tropical to subarctic zones. Can J Zool 72: 1462–1468.
- 143.↑
Mogi M , 1981. Studies on Aedes togoi (Diptera: Culicidae): 1. Alternative diapause in the Nagasaki strain. J Med Entomol 18: 477–480.
- 144.↑
Galka BE , Brust RA , 1987. The effect of temperature and photoperiod on the induction of larval diapause in the mosquito Aedes togoi (Theobald) (Diptera: Culicidae). Can J Zool 65: 2262–2265.
- 145.↑
Sollers-Riedel H , 1971. World studies on mosquitoes and diseases carried by them. Proceedings of the 58th Annual Meeting of the New Jersey Mosquito Extermination Association, Atlantic City. NJ, Supplement, 1–52.
- 146.↑
Trimble R , Wellington W , 1979. Colonization of North American Aedes togoi. Mosq News 39: 18–20.
- 149.↑
Rosen L , Tesh R , Lien J , Cross J , 1978. Transovarial transmission of Japanese encephalitis virus by mosquitoes. Science 199: 909–911.
- 150.↑
Rosen L , 1986. The natural history of Japanese encephalitis virus. Annu Rev Microbiol 40: 395–414.
- 151.↑
Becker N , Petric D , Zgomba M , Boase C , Minoo M , Dahl C , Kaiser A , 2010. Mosquitoes and Their Control, 2nd edition. Heidelberg, Germany: Springer.
- 152.↑
Tsuda Y , Komagata O , Kasai S , Hayashi T , Nihei N , Saito K , Mizutani M , Kunida M , Yoshida M , Kobayashi M , 2008. A mark–release–recapture study on dispersal and flight distance of Culex pipiens pallens in an urban area of Japan. J Am Mosq Control Assoc 24: 339–343.
- 153.↑
Fritz ML , Walker ED , Yunker AJ , Dworkin I , 2014. Daily blood feeding rhythms of laboratory-reared North American Culex pipiens. J Circadian Rhythms 12: 1.
- 154.↑
Peach DAH , Gries G , 2020. Mosquito phytophagy—sources exploited, ecological function, and evolutionary transition to haematophagy. Entomol Exp Appl 168: 120–136.
- 155.↑
Peach DAH , Gries G , 2016. Nectar thieves or invited pollinators? A case study of tansy flowers and common house mosquitoes. Arthropod-Plant Interact 10: 497–506.
- 156.↑
Onyeka JOA , Boreham PFL , 1987. Population studies, physiological state and mortality factors of overwintering adult populations of females of Culex pipiens L. (Diptera: Culicidae). Bull Entomol Res 77: 99–111.
- 157.↑
Bailey CL , Faran ME , Gargan TP , Hayes DE , 1982. Winter survival of blood-fed and nonblood-fed Culex pipiens L. Am J Trop Med Hyg 31: 1054–1061.
- 158.↑
Giberson DJ , Dau-Schmidt K , Dobrin M , 2007. Mosquito species composition, phenology and distribution (Diptera: Culicidae) on Prince Edward Island. Entomol Soc J Acadian Entomol Soc 3: 7–27.
- 159.↑
Chaulk AC , Carson KP , Whitney HG , Fonseca DM , Chapman TW , 2016. The arrival of the northern house mosquito Culex pipiens (Diptera: Culicidae) on Newfoundland’s Avalon Peninsula. J Med Entomol 53: 1364–1369.
- 160.↑
Hongoh V , Berrang-Ford L , Scott ME , Lindsay LR , 2012. Expanding geographical distribution of the mosquito, Culex pipiens, in Canada under climate change. Appl Geogr 33: 53–62.
- 161.↑
Harbach RE , 2012. Culex pipiens: species versus species complex—taxonomic history and perspective. J Am Mosq Control Assoc 28: 10–23.
- 163.↑
Hearle E , 1926. The Mosquitoes of the Lower Fraser Valley, British Columbia, and their control. Ottawa, ON: Natural Research Council of Canada Report 17, 94.
- 164.↑
Peach DAH , Poirier LM. , 2020. New distribution records and range extensions of mosquitoes in British Columbia and the Yukon Territory. J Entomol Soc Br Colum Subm117: 69–74.
- 165.↑
Coquillet DW , 1900. Papers from the Harriman Alaska expedition. IX. Entomological results (3): Diptera. Proc Wash Acad Sci 2: 389–464.
- 166.↑
Russell C , Hunter FF , 2012. Culex pipiens (Culicidae) is attracted to humans in southern Ontario, but will it serve as a bridge vector of West Nile virus? Can Entomol 144: 667–671.
- 167.↑
Francy DB , Rush WA , Montoya M , Inglish DS , Bolin RA , 1981. Transovarial transmission of St. Louis encephalitis virus by Culex pipiens complex mosquitoes. Am J Trop Med Hyg 30: 699–705.
- 168.↑
Mitchell CJ , Bowen GS , Monath TP , Cropp CB , Kerschner J , 1979. St. Louis encephalitis virus transmission following multiple feeding of Culex pipiens pipiens (Diptera: Culicidae) during a single gonotrophic cycle. J Med Entomol 16: 254–258.
- 169.↑
Camp JV , Kolodziejek J , Nowotny N , 2019. Targeted surveillance reveals native and invasive mosquito species infected with Usutu virus. Parasit Vectors 12: 46.
- 170.↑
Glaizot O , Fumagalli L , Iritano K , Lalubin F , Van Rooyen J , Christe P , 2012. High prevalence and lineage diversity of avian malaria in wild populations of great tits (Parus major) and mosquitoes (Culex pipiens). PLoS One 7: e34964.
- 171.↑
Lalubin F , Delédevant A , Glaizot O , Christe P , 2013. Temporal changes in mosquito abundance (Culex pipiens), avian malaria prevalence and lineage composition. Parasit Vectors 6: 307.
- 172.↑
Schoener ER , Harl J , Himmel T , Fragner K , Weissenböck H , Fuehrer H-P , 2019. Protozoan parasites in Culex pipiens mosquitoes in Vienna. Parasitol Res 118: 1261–1269.
- 173.↑
Spence Beaulieu MR , Federico JL , Reiskind MH , 2020. Mosquito diversity and dog heartworm prevalence in suburban areas. Parasit Vectors 13: 12.
- 174.↑
Dadum D , Robinson R , Clements A , Peach W , Bennett M , Rowcliffe JM , Cunningham A , 2019. Avian malaria-mediated population decline of a widespread iconic bird species. R Soc Open Sci 6: 182197.
- 175.↑
Atkinson CT , Woods KL , Dusek RJ , Sileo LS , Iko WM , 1995. Wildlife disease and conservation in Hawaii: pathogenicity of avian malaria (Plasmodium relictum) in experimentally infected Iiwi (Vestiaria coccinea). Parasitology 111: S59–S69.
- 176.↑
van Riper C , van Riper SG , Goff ML , Laird M , 1986. The epizootiology and ecological significance of malaria in Hawaiian land birds. Ecol Monogr 56: 327–344.
- 177.↑
Scott JM , Mountainspring S , Ramsey FL , Kepler CB , 1989. Forest bird communities of the Hawaiian Islands: their dynamics, ecology, and conservation. J Wildl Manage 53: 1–431.
- 178.↑
Vanderwerf EA , Burt MD , Rohrer JL , Mosher SM , 2006. Distribution and prevalence of mosquito-borne diseases in O’ahu ’Elepaio. Condor 108: 770–777.
- 179.↑
Restifo RA , 1980. The occurrence of Aedes atropalpus (Coquitllett) breeding in tires in Ohio and Indiana. Mosq News 40: 292–294.
- 180.↑
Romi R , Sabatinelli G , Savelli LG , Raris M , Zago M , Malatesta R , 1997. Identification of a North American mosquito species, Aedes atropalpus (Diptera: Culicidae), in Italy. J Am Mosq Control Assoc 13: 245–246.
- 181.↑
Scholte EJ , Beeuwkes J , 2010. Introduction and control of three invasive mosquito species in the Netherlands, July–October 2010. Euro Surveill 15: 1–4.
- 182.↑
Montarsi F et al.2013. Distribution and habitat characterization of the recently introduced invasive mosquito Aedes koreicus [Hulecoeteomyia koreica], a new potential vector and pest in north-eastern Italy. Parasit Vectors 6: 292.
- 183.↑
Werner D , Zielke DE , Kampen H , 2016. First record of Aedes koreicus (Diptera: Culicidae) in Germany. Parasitol Res 115: 1331–1334.
- 184.↑
Asahina S , 1970. Transoceanic flight of mosquitoes on the Northwest Pacific. JJMSB 23: 255–258.
- 185.↑
Sunahara T , Ishizaka K , Mogi M , 2002. Habitat size: a factor determining the opportunity for encounters between mosquito larvae and aquatic predators. J Vector Ecol 13: 8–20.
- 186.↑
Seebens H et al.2017. No saturation in the accumulation of alien species worldwide. Nat Commun 8: 14435.
- 187.↑
Hebert PDN , Cywinska A , Ball SL , deWaard JR , 2003. Biological identifications through DNA barcodes. Proc Biol Sci 270: 313–321.
- 188.↑
Dhananjeyan KJ , Paramasivan R , Tewari SC , Rajendran R , Thenmozhi V , Leo SVJ , Venkatesh A , Tyagi BK , 2010. Molecular identification of mosquito vectors using genomic DNA isolated from eggshells, larval and pupal exuvium. Trop Biomed 27: 47–53.
- 189.↑
Jackson M , Pyles C , Breton S , McMahon T , Belton P , 2013. British Columbia’s 50th mosquito species, Aedes schizopinax. J Entomol Soc BC 110: 38–39.
- 190.↑
Brown JE , Evans BR , Zheng W , Obas V , Barrera-Martinez L , Egizi A , Zhao H , Caccone A , Powell JR , 2014. Human impacts have shaped historical and recent evolution in Aedes aegypti, the dengue and yellow fever mosquito. Evolution 68: 514–525.
- 191.↑
Brown JE , Scholte E-J , Dik M , Den Hartog W , Beeuwkes J , Powell JR , 2011. Aedes aegypti mosquitoes imported into the Netherlands, 2010. Emerg Infect Dis 17: 2335–2337.
- 192.↑
Logue K , Keven JB , Cannon MV , Reimer L , Siba P , Walker ED , Zimmerman PA , Serre D , 2016. Unbiased characterization of Anopheles mosquito blood meals by targeted high-throughput sequencing. PLoS Negl Trop Dis 10: e0004512.
- 193.↑
Kent RJ , 2009. Molecular methods for arthropod bloodmeal identification and applications to ecological and vector-borne disease studies. Mol Ecol Resour 9: 4–18.
- 194.↑
Crabtree MB , Kading RC , Mutebi J-P , Lutwama JJ , Miller BR , 2013. Identification of host blood from engorged mosquitoes collected in western Uganda using cytochrome oxidase I gene sequences. J Wildl Dis 49: 611–626.
- 195.↑
Ngo KA , Kramer LD , 2003. Identification of mosquito bloodmeals using polymerase chain reaction (PCR) with order-specific primers. J Med Entomol 40: 215–222.
- 196.↑
Reeves LE , Gillett-Kaufman JL , Kawahara AY , Kaufman PE , 2018. Barcoding blood meals: new vertebrate-specific primer sets for assigning taxonomic identities to host DNA from mosquito blood meals. PLoS Negl Trop Dis 12: e0006767.
- 197.↑
Gyawali N , Taylor-Robinson AW , Bradbury RS , Huggins DW , Hugo LE , Lowry K , Aaskov JG , 2019. Identification of the source of blood meals in mosquitoes collected from north-eastern Australia. Parasit Vectors 12: 198.
- 198.↑
Harrington LC et al.2014. Heterogeneous feeding patterns of the dengue vector, Aedes aegypti, on individual human hosts in rural Thailand. PLoS Negl Trop Dis 8: e3048.
- 199.↑
Kingan SB , Heaton H , Cudini J , Lambert CC , Baybayan P , Galvin BD , Durbin R , Korlach J , Lawniczak MKN , 2019. A high-quality de novo genome assembly from a single mosquito using PacBio sequencing. Genes (Basel) 10: 62.
- 200.↑
Matthews BJ et al.2018. Improved reference genome of Aedes aegypti informs arbovirus vector control. Nature 563: 501–507.
- 201.↑
Ghurye J , Koren S , Small ST , Redmond S , Howell P , Phillippy AM , Besansky NJ , 2019. A chromosome-scale assembly of the major African malaria vector Anopheles funestus. Gigascience 8: giz063.
- 202.↑
Kothera L , Phan J , Ghallab E , Delorey M , Clark R , Savage HM , 2019. Using targeted next-generation sequencing to characterize genetic differences associated with insecticide resistance in Culex quinquefasciatus populations from the southern U.S. PLoS One 14: e0218397.
- 203.↑
Liu N , 2015. Insecticide resistance in mosquitoes: impact, mechanisms, and research directions. Annu Rev Entomol 60: 537–559.
- 204.↑
Demok S , Endersby-Harshman N , Vinit R , Timinao L , Robinson LJ , Susapu M , Makita L , Laman M , Hoffmann A , Karl S , 2019. Insecticide resistance status of Aedes aegypti and Aedes albopictus mosquitoes in Papua New Guinea. Parasit Vectors 12: 333.
- 205.↑
Faucon F et al.2015. Identifying genomic changes associated with insecticide resistance in the dengue mosquito Aedes aegypti by deep targeted sequencing. Genome Res 25: 1347–1359.
- 206.↑
Rose NH et al.2020. Climate and urbanization drive mosquito preference for humans. Curr Biol 30: 3570–3579.
- 207.↑
Connelly R , 2019. Highlights of medical entomology 2018: the importance of sustainable surveillance of vectors and vector-borne pathogens. J Med Entomol 56: 1183–1187.
- 208.↑
Borkent A , 2018. The state of phylogenetic analysis: narrow visions and simple answers—examples from the Diptera (flies). Zootaxa 4374: 107.
- 209.↑
Piper AM , Batovska J , Cogan NOI , Weiss J , Cunningham JP , Rodoni BC , Blacket MJ , 2019. Prospects and challenges of implementing DNA metabarcoding for high-throughput insect surveillance. Gigascience 8: giz092.
- 210.↑
Parker C , Ramirez D , Connelly CR , 2019. State‐wide survey of Aedes aegypti and Aedes albopictus (Diptera: Culicidae) in Florida. J Vector Ecol 44: 210–215.
- 211.↑
Jourdain F et al.2019. Towards harmonisation of entomological surveillance in the Mediterranean area. PLoS Negl Trop Dis 13: e0007314.
- 212.↑
World Health Organization , 2017. Global Vector Control Response 2017–2030. Geneva, Switzerland: WHO.
- 213.↑
Ammar SE , Mclntyre M , Swan T , Kasper J , Derraik JGB , Baker MG , Hales S , 2019. Intercepted mosquitoes at New Zealand’s ports of entry, 2001 to 2018: Current status and future concerns. TropicalMed 4: 101.
- 214.↑
Medlock JM , Vaux AG , Cull B , Schaffner F , Gillingham E , Pfluger V , Leach S , 2017. Detection of the invasive mosquito species Aedes albopictus in southern England. Lancet Infect Dis 17: 140.
- 215.↑
Ibañez-Justicia A , Gloria-Soria A , den Hartog W , Dik M , Jacobs F , Stroo A , 2017. The first detected airline introductions of yellow fever mosquitoes (Aedes aegypti) to Europe, at Schiphol International airport, the Netherlands. Parasit Vectors 10: 603.
- 216.↑
Sukehiro N , Kida N , Umezawa M , Murakami T , Arai N , Jinnai T , Inagaki S , Tsuchiya H , Maruyama H , Tsuda Y , 2013. First report on invasion of yellow fever mosquito, Aedes aegypti, at Narita International Airport, Japan in August 2012. Jpn J Infect Dis 66: 189–194.
- 217.↑
Kampen H , Zielke D , Werner D , 2012. A new focus of Aedes japonicus japonicus (Theobald, 1901) (Diptera, Culicidae) distribution in Western Germany: rapid spread or a further introduction event? Parasit Vectors 5: 284.
- 218.↑
Kaplan L , Kendell D , Robertson D , Livdahl T , Khatchikian C , 2010. Aedes aegypti and Aedes albopictus in Bermuda: extinction, invasion, invasion and extinction. Biol Invasions 12: 3277–3288.
- 219.↑
Thielman A , 2011. Canadian National Collection of Insects, Arachnids and Nematodes - Culicidae Database. Available at: http://canacoll.org/Diptera/Aynsley/index.htm. Accessed January 2019.
- 220.↑
Ontario Agency for Health Protection and Promotion (Public Health Ontario) , 2020. WNV Mosquito ID Data for Ochlerotatus japonicus From 2002–2019. Toronto, Ontario: Public Health Ontario.