Aedes triseriatus

Ge­o­graphic Range

East­ern tree­hole mos­qui­toes (Aedes tris­e­riatus) are the most com­mon tree­hole mos­qui­toes in the United States. Na­tive to the Nearc­tic re­gion, they can be found in south­ern Canada and the east­ern United States. Their range ex­tends as far south as the Florida Keys, and west to Idaho, Utah, and Texas. In Canada, they are wide­spread in the south­ern parts of British Co­lum­bia, Que­bec, On­tario, and Saskatchewan. While they have not es­tab­lished pop­u­la­tions over­seas, east­ern tree­hole mos­qui­toes do have the po­ten­tial for range ex­pan­sion. In 2001, a spec­i­men was found in France in a ship­ment of used tires from the United States, but was im­me­di­ately killed. With the preva­lence of over­seas travel and com­merce, east­ern tree­hole mos­qui­toes could cer­tainly es­tab­lish them­selves in re­gions of Eu­rope, if care is not taken. (Car­pen­ter and La Casse, 1955; Fara­jol­lahi and Price, 2013; Med­lock, et al., 2012)

Habi­tat

Lar­vae of east­ern tree­hole mos­qui­toes are aquatic, and live in wa­ter-filled cav­i­ties, typ­i­cally in de­cid­u­ous trees, giv­ing the species their com­mon name. Lar­vae can also be found in man-made con­tain­ers that col­lect rain water, such as tubs, bar­rels, and aban­doned tires. Adult fe­males tend to oviposit in forested, shaded areas with dense under sto­ries, gen­er­ally avoid­ing open areas. Adults are ter­res­trial, and are usu­ally found in forests, as well as sub­ur­ban areas lo­cated near forests. (Car­pen­ter and La Casse, 1955; Ellis, 2008; Leisnham and Ju­liano, 2012)

Phys­i­cal De­scrip­tion

East­ern tree­hole mos­qui­toes are a medium-sized mos­quito species. Fe­males are gen­er­ally larger, al­though the col­oration is sim­i­lar be­tween the sexes. Their pro­boscis and their short palpi are black. The pos­te­rior of their head is cov­ered in sil­ver-white scales. Their tho­rax is brown or black, with a wide me­dian stripe of dark brown, and their sides are white. Their scutel­lum is dark brown. Their ab­domen is blue-black, with patches of white lat­er­ally. Their wings are about 3.5 to 4.0 mm in length, with dark scales. Their hind fe­murs are yel­low­ish-white, and dark to­wards the ends. Femora, tibia, and tarsi are all black. Lar­vae are long and thin, with seg­mented bod­ies, and are typ­i­cally cream col­ored or brown. They have a breath­ing tube on their pos­te­rior end. East­ern tree­hole mos­quito lar­vae can be dis­tin­guished from re­lated species by comb scales in dou­ble roles, anal papilla of un­equal length, and multi-branched lat­eral hair on the anal sad­dle. Pupae are a sim­i­lar color, with a shorter, curved body. (Car­pen­ter and La Casse, 1955; Costanzo, et al., 2011; Fara­jol­lahi and Price, 2013)

  • Sexual Dimorphism
  • female larger

De­vel­op­ment

East­ern tree­hole mos­qui­toes are holometabolous. Eggs over­win­ter on the sides of tree holes and other con­tain­ers, and hatch in the spring when they are cov­ered by water col­lected in the cav­i­ties, in re­sponse to the lack of oxy­gen. Water tem­per­a­tures must be 4.1 to 12 de­grees Cel­sius for eggs to begin hatch­ing. Hatch­ing is also in­flu­enced by a va­ri­ety of other en­vi­ron­men­tal fac­tors, and the eggs usu­ally do not all hatch at once. Hatch­ings occur in waves, which is use­ful in case of drought. If the tree hole dries up, the lar­vae will die, but eggs that have not yet hatched are still vi­able for when the tree hole fills with water again. Some eggs hatch after the first rain­fall, while oth­ers do not hatch until there have been many rain­falls. Hatch­ing can also be de­layed if there is a high lar­val den­sity al­ready in the tree hole, or in re­sponse to older mos­quito lar­vae in the water, which al­lows for newly hatched lar­vae to avoid com­pet­ing for re­sources with older in­stars. Once the eggs hatch, there are four lar­val in­stars. Time of lar­val de­vel­op­ment de­pends on re­source avail­abil­ity, with higher amounts of de­tri­tus al­low­ing for faster de­vel­op­ment and larger adults. They then molt into pupae, and then eclose and leave the water as adults. (Car­pen­ter and La Casse, 1955; Edgerly and Liv­dahl, 1992; Ellis, 2008; Har­shaw, et al., 2007; Khatchikian, et al., 2009; Leisnham and Ju­liano, 2012; Williams, et al., 2007)

The de­vel­op­ment of east­ern tree­hole mos­qui­toes has a high level of plas­tic­ity, and varies among re­gions and even from year to year. This is also due to de­layed hatch­ings, amount of rain­fall, tem­per­a­ture, and re­source avail­abil­ity. As a re­sult, there are usu­ally mul­ti­ple co­horts and all in­star stages pre­sent at once. Some pop­u­la­tions have been ob­served as uni­vol­tine, with one pop­u­la­tion per year. In these pop­u­la­tions, lar­vae first ap­peared in mid-March, 2nd in­stars ap­peared in mid-April, with many first in­stars pre­sent. Four weeks later, all in­star stages were pre­sent, with some pupae, and then adults pre­sent in June and July. Other pop­u­la­tions have mul­ti­ple gen­er­a­tions per year. First in­stars first ap­peared in mid-April, and after 3 weeks, some 4th in­star lar­vae were pre­sent. Pupae were pre­sent by the end of May. The first gen­er­a­tion took about 6 weeks to de­velop into adults. In early June, a sec­ond gen­er­a­tion oc­curred, re­sult­ing in pupae 2 to 3 weeks later. A third gen­er­a­tion de­vel­oped into 2nd in­star lar­vae by early June, pu­pat­ing within a week or two af­ter­ward. De­vel­op­ment of the later gen­er­a­tions took only about 3 weeks. Even small fourth gen­er­a­tions have been ob­served, be­gin­ning in mid-Au­gust, though these might not have enough time to de­velop into adults be­fore win­ter ar­rives. In some mid­dle re­gions of their range, where the tree holes do not com­pletely freeze, some lar­vae will di­a­pause dur­ing the win­ter, in ad­di­tion to over­win­ter­ing eggs. In the most south­ern parts of their range, east­ern tree­hole mos­qui­toes are ac­tive year round. (Car­pen­ter and La Casse, 1955; Edgerly and Liv­dahl, 1992; Ellis, 2008; Har­shaw, et al., 2007; Khatchikian, et al., 2009; Leisnham and Ju­liano, 2012; Williams, et al., 2007)

Re­pro­duc­tion

Fe­male east­ern tree­hole mos­qui­toes must take a blood meal be­fore mat­ing, to ma­ture a batch of eggs. Males find fe­male mates by de­tect­ing the buzzing sound made by their beat­ing wings, which is species spe­cific. Mat­ing typ­i­cally oc­curs in the mid­dle of the sum­mer, from June to July, but due to the high plas­tic­ity of this species' de­vel­op­ment cycle, mat­ing may also occur later in the sea­son as later gen­er­a­tions de­velop quickly and emerge as adults. In the south­ern­most re­gion of their range, they mate year round. Typ­i­cally, fe­males only mate once, but some fe­males live long enough to take a sec­ond blood meal and un­dergo a sec­ond ovipo­si­tion cycle. In very rare cir­cum­stances, fe­males live long enough to un­dergo a third cycle. Hy­bridiza­tions have been recorded oc­cur­ring be­tween east­ern tree­hole mos­qui­toes and other mos­quito species, in­clud­ing Aedes hen­der­soni. In­ter­est­ingly, fe­males in­fected with La Crosse en­cephali­tis virus, for which this species is a vec­tor, are more ef­fi­cient at mat­ing than non-in­fected fe­males. In­fected fe­males have an in­creased, ear­lier sperm trans­fer. (Fara­jol­lahi and Price, 2013; Fara­jol­lahi and Price, 2013; Frank­ino and Ju­liano, 1999; Gib­son and Rus­sell, 2006; Reese, et al., 2009; Spiel­man and D'An­to­nio, 2001)

After mat­ing, eggs are laid on the side of water filled holes or other man-made con­tain­ers. The eggs are laid singly or in groups of 2 to 5. They are laid just above the water line, and will not hatch until they have been cov­ered with water of a cer­tain tem­per­a­ture, along with sev­eral other fac­tors. Fe­males typ­i­cally only lay one batch of eggs, though some do sur­vive long enough to mate a sec­ond time and lay a sec­ond batch. Stud­ies have shown that fe­males are more likely to lay eggs in cav­i­ties that al­ready have eggs in them. The pres­ence of or­ganic mat­ter in the water is also pos­si­bly an ap­peal­ing fac­tor for fe­males when search­ing for an ovipo­si­tion site. Stud­ies have also shown fe­male east­ern tree­hole mos­qui­toes are more at­tracted to dyed water. Ovipo­si­tion rates are re­port­edly high­est in July, though this varies by re­gion, as the south­ern­most pop­u­la­tions can breed year round. La Crosse en­cephali­tis virus is trans­mit­ted ver­ti­cally from par­ent to off­spring. The virus over­win­ters in the eggs. (Beehler, et al., 1992; Car­pen­ter and La Casse, 1955; Ellis, 2008; Reese, et al., 2009; Spiel­man and D'An­to­nio, 2001)

  • Breeding interval
    Eastern treehole mosquitoes typically mate once, though some females survive for multiple mating and egg laying cycles.
  • Breeding season
    Breeding occurs during June and July, but this varies by region.

Adult east­ern tree­hole mos­qui­toes pro­vide pro­vi­sion­ing in the eggs, as well as lay the eggs in suit­able wa­ter-filled tree holes or other cav­i­ties. Oth­er­wise, there is no more parental care. (Car­pen­ter and La Casse, 1955)

  • Parental Investment
  • pre-hatching/birth
    • provisioning
      • female

Lifes­pan/Longevity

Fe­males live any­where from about 2 to 5 weeks after reach­ing adult­hood. Males likely live for a shorter pe­riod of time. (Frank­ino and Ju­liano, 1999)

  • Typical lifespan
    Status: wild
    11 to 40 days
  • Typical lifespan
    Status: captivity
    11 to 40 days

Be­hav­ior

Adult east­ern tree­hole mos­qui­toes are mainly cre­pus­cu­lar, with the fe­males tak­ing most of their blood meals dur­ing dawn and dusk. Males are ac­tive dur­ing this time, for­ag­ing for food, and search­ing for mates. Fe­males in­fected with La Crosse virus show be­hav­ioral changes. When tak­ing blood meals, in­fected fe­males probe more and en­gorge less. This re­sults in more feed­ings, re­sult­ing in more virus trans­mis­sions. Lar­vae can often be found float­ing near the water sur­face, with their breath­ing tubes stick­ing out of the water. Lar­vae are rea­son­ably ac­tive, and move with a wrig­gling mo­tion. They leave the sur­face to feed in the ben­thos and water col­umn, and dart down to the bot­tom of the cav­ity when dis­turbed. Lar­val den­sity can be very high in their tree holes. Stud­ies have found any­where from 60 lar­vae per 100 ml to 150 or more per 100 ml. Pupae tend to be much less ac­tive than lar­vae, and re­main float­ing at the sur­face of the water. They can still sense a threat, how­ever, and can move away when nec­es­sary. (Car­pen­ter and La Casse, 1955; Reese, et al., 2009; Williams, et al., 2007)

Home Range

Lar­val live in small tree holes and con­tain­ers, which they do not leave until eclo­sion. Adults likely re­main in the same gen­eral re­gion from which they emerged. (Car­pen­ter and La Casse, 1955)

Com­mu­ni­ca­tion and Per­cep­tion

The pri­mary sen­sory struc­tures for mos­qui­toes are their an­ten­nae, pro­boscis, and tip of their ab­domen. Mos­qui­toes ac­tively groom these sen­silla, likely clear­ing them of ob­struc­tive par­ti­cles. At the base of the an­ten­nae is an au­di­tory organ. Males iden­tify mates though sound, by de­tect­ing the wing beat­ing tones of the fe­males, which cre­ates the buzzing sound that mos­qui­toes are known for pro­duc­ing in flight. This sound is species spe­cific. When search­ing for blood meals, fe­males typ­i­cally de­tect chem­i­cals and other cues to find hosts, such as car­bon diox­ide. They can also vi­su­ally de­tect hosts, other mos­qui­toes, and the en­vi­ron­ment as a whole. Fe­males use their pro­boscis to tac­tilely probe the skin of the host, and may rein­sert the pro­boscis sev­eral times until they have found a suit­able blood ves­sel. Light is also typ­i­cally a strong at­trac­tant for mos­qui­toes, but some stud­ies have re­ported that light traps are not par­tic­u­larly ef­fec­tive for catch­ing east­ern tree­hole mos­qui­toes. Lar­vae can also de­tect chem­i­cal sig­nals, typ­i­cally alarm sig­nals, which would be pro­duced by a neigh­bor­ing larva when at­tacked or eaten by a preda­tor. (Costanzo, et al., 2011; Gib­son and Rus­sell, 2006; Spiel­man and D'An­to­nio, 2001; Walker and Archer, 1988)

Food Habits

Adult mos­qui­toes are typ­i­cally nec­tari­vores. Males feed solely on nec­tar, while fe­males take blood meals in ad­di­tion to nec­tar. These acts of par­a­sitism must occur be­fore a fe­male is able to ma­ture her eggs and mate. Small mam­mals are the typ­i­cal host of east­ern tree­hole mos­qui­toes, par­tic­u­larly chip­munks and squir­rels. They have also been ob­served feed­ing from birds, as well as larger mam­mals, in­clud­ing hu­mans. Lar­vae of east­ern tree­hole mos­qui­toes are de­tri­ti­vores. They fil­ter feed and browse or­ganic de­tri­tus from leaf lit­ter, as well as mi­crobes and par­ti­cles of de­cay­ing in­ver­te­brates. Their feed­ing takes place pri­mar­ily in the ben­thos and water col­umn. Though there is some dis­agree­ment be­tween re­searchers, can­ni­bal­ism of ear­lier in­stars by older lar­vae may occur. (Car­pen­ter and La Casse, 1955; Ellis, 2008; Har­shaw, et al., 2007; Koenekoop and Liv­dahl, 1986; Spiel­man and D'An­to­nio, 2001; Tuten, et al., 2012)

  • Animal Foods
  • blood

Pre­da­tion

The main preda­tors of east­ern tree­hole mos­quito lar­vae are other lar­vae that co-habi­tate in their wa­ter-filled tree holes and cav­i­ties. Lar­vae of the mos­qui­toes Tox­orhyn­chites ru­tilus and Anophe­les bar­beri, as well as lar­vae of the midge Corethrella ap­pen­dic­u­lata, are preda­tory and are often found in tree holes with east­ern tree­hole mos­qui­toes. In some re­gions of the coun­try, the lar­vae face lit­tle to no pre­da­tion. In the pres­ence of a preda­tor, east­ern tree­hole mos­quito lar­vae de­crease their ac­tiv­ity, often by just rest­ing at the sur­face. Preda­tors of adults in­clude ants, bee­tles, preda­tory hemipter­ans, bats, birds, and other op­por­tunis­tic ver­te­brates. Adults are es­pe­cially vul­ner­a­ble to ter­res­trial preda­tors shortly after eclo­sion, when they are weak and un­steady. (Alto, et al., 2009; Costanzo, et al., 2011; Ellis, 2008; Nan­nini and Ju­liano, 1998; Rochlin, et al., 2013; Spiel­man and D'An­to­nio, 2001)

Ecosys­tem Roles

East­ern tree­hole mos­quito lar­vae have a sig­nif­i­cant pres­ence in tree hole com­mu­ni­ties, and are often the dom­i­nant species. They are often found in the same wa­ter-filled tree holes and cav­i­ties as other lar­val mos­quito species, in­clud­ing Anophe­les bar­beri, Or­thopodomyia species, Tox­orhyn­chites ru­tilus septen­tri­on­alis, and Aedes zooso­phus. East­ern tree­hole mos­qui­toes also occur sym­patri­cally with Aedes hen­der­soni, and hy­bridiza­tions can occur be­tween the two species. Lar­vae are often prey to preda­tory mos­quito lar­vae in­hab­it­ing the same tree holes. East­ern tree­hole mos­quito lar­vae can also be in­fected by par­a­sitic gre­garines, As­cogre­ga­rina bar­retti, which are found in the gut of the lar­vae, as well as in pupae and adults. There has been much focus on the in­ter­ac­tions of east­ern tree­hole mos­qui­toes with other in­va­sive tree hole mos­qui­toes. Lar­vae of Aedes al­bopic­tus, the Asian tiger mos­quito, and Aedes japon­i­cus, often live in the same tree holes. Aedes al­bopic­tus and A. japon­cius seem to out­com­pete east­ern tree­hole mos­qui­toes for re­sources, which has re­sulted in some de­clines in their pop­u­la­tion. Since Aedes al­bopic­tus can also vec­tor the La Crosse en­cephali­tis virus, this is un­for­tu­nately not help­ful to hu­mans and other virus hosts. How­ever, east­ern tree­hole mos­qui­toes seem to be able to still sur­vive rea­son­ably well in the same tree holes with these in­va­sive species. Aedes al­bopic­tus ap­pears to be more sus­cep­ti­ble to pre­da­tion and east­ern tree­hole mos­qui­toes also over­win­ter more suc­cess­fully as eggs, and emerges first, al­low­ing them to col­o­nize tree holes and uti­lize re­sources ear­lier than ei­ther of the in­va­sive species, al­low­ing them to co-ex­ist. (Alto, et al., 2009; Car­pen­ter and La Casse, 1955; Fara­jol­lahi and Price, 2013; Leisnham and Ju­liano, 2012; Rochlin, et al., 2013)

Fe­male east­ern tree­hole mos­qui­toes are par­a­sitic, and re­quire blood meals be­fore they can suc­cess­fully re­pro­duce. They typ­i­cally feed on small mam­mals, such as chip­munks and squir­rels, though they may also feed on birds and other larger mam­mals, in­clud­ing hu­mans. East­ern tree­hole mos­qui­toes are the main vec­tors for the La Crosse en­cephali­tis virus in the United States, and can trans­mit the dis­ease to hu­mans, as well as east­ern chip­munks, gray squir­rels, and red foxes. They also trans­mit a large va­ri­ety of other dis­eases to hu­mans and many other do­mes­tic and non-do­mes­tic an­i­mals, in­clud­ing West Nile virus, east­ern equine en­cephali­tis, and Diro­fi­laria im­mi­tis. (Fara­jol­lahi and Price, 2013; Leisnham and Ju­liano, 2012; Reese, et al., 2009; Tuten, et al., 2012)

Species Used as Host
Com­men­sal/Par­a­sitic Species
  • gre­garines (As­cogre­ga­rina bar­retti)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

East­ern tree­hole mos­qui­toes have pro­vided a vast amount of re­search op­por­tu­ni­ties, as sci­en­tists study the tree hole com­mu­ni­ties, their in­ter­ac­tions with and re­sponses to in­va­sive mos­quito species, and their role as a vec­tor for a large va­ri­ety of both human and zoonotic dis­eases. Oth­er­wise, east­ern tree­hole mos­qui­toes have no pos­i­tive ef­fects on hu­mans. (Fara­jol­lahi and Price, 2013)

  • Positive Impacts
  • research and education

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Fe­male east­ern tree­hole mos­qui­toes are par­a­sitic, and take blood meals from many small mam­mals, as well as hu­mans. Their bites can be painful and very ir­ri­tat­ing. East­ern tree­hole mos­qui­toes are the main vec­tors of the La Crosse en­cephali­tis virus in the United States. La Crosse en­cephali­tis often has no symp­toms, but can cause fever, headache, nau­sea, vom­it­ing, and fa­tigue. Se­vere cases, which occur most often in chil­dren, can re­sult in se­vere neu­roin­va­sive dis­ease, as well as en­cephali­tis, which can cause seizures, coma, and paral­y­sis. Long term dis­abil­ity or death can occur in very rare cases. There are about 80 to 100 cases in the United States each year, but this num­ber is likely un­der­re­ported, as La Crosse virus can have no symp­toms. In ad­di­tion to La Crosse virus, east­ern tree­hole mos­qui­toes can also vec­tor Cache Val­ley virus, east­ern equine en­cephali­tis, High­lands J virus, Jamestown Canyon virus, and West Nile virus. Ad­di­tion­ally, they can also trans­mit Diro­fi­laria im­mi­tis, which causes heart­worm in dogs, cats, and other an­i­mals. (CDC, 2009; Car­pen­ter and La Casse, 1955; Fara­jol­lahi and Price, 2013)

Con­ser­va­tion Sta­tus

East­ern tree­hole mos­qui­toes have no spe­cial con­ser­va­tion sta­tus. Since they can trans­mit many dis­eases to hu­mans and other an­i­mals, ef­forts are di­rected to­wards con­trol rather than con­ser­va­tion.

Other Com­ments

Aedes tris­e­riatus is also known as Ochlero­ta­tus tris­e­riatus. (Fara­jol­lahi and Price, 2013)

Con­trib­u­tors

An­gela Miner (au­thor), An­i­mal Di­ver­sity Web Staff, Leila Si­cil­iano Mar­tina (ed­i­tor), An­i­mal Di­ver­sity Web Staff.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

acoustic

uses sound to communicate

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

causes or carries domestic animal disease

either directly causes, or indirectly transmits, a disease to a domestic animal

chemical

uses smells or other chemicals to communicate

crepuscular

active at dawn and dusk

detritus

particles of organic material from dead and decomposing organisms. Detritus is the result of the activity of decomposers (organisms that decompose organic material).

diapause

a period of time when growth or development is suspended in insects and other invertebrates, it can usually only be ended the appropriate environmental stimulus.

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

fertilization

union of egg and spermatozoan

filter-feeding

a method of feeding where small food particles are filtered from the surrounding water by various mechanisms. Used mainly by aquatic invertebrates, especially plankton, but also by baleen whales.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

freshwater

mainly lives in water that is not salty.

herbivore

An animal that eats mainly plants or parts of plants.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

internal fertilization

fertilization takes place within the female's body

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nectarivore

an animal that mainly eats nectar from flowers

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

parasite

an organism that obtains nutrients from other organisms in a harmful way that doesn't cause immediate death

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

semelparous

offspring are all produced in a single group (litter, clutch, etc.), after which the parent usually dies. Semelparous organisms often only live through a single season/year (or other periodic change in conditions) but may live for many seasons. In both cases reproduction occurs as a single investment of energy in offspring, with no future chance for investment in reproduction.

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

visual

uses sight to communicate

Ref­er­ences

Alto, B., B. Ke­savaraju, S. Ju­liano, L. Louni­bos. 2009. Stage-de­pen­dent pre­da­tion on com­peti­tors: con­se­quences for the out­come of a mos­quito in­va­sion. Jour­nal of An­i­mal Ecol­ogy, 78/5: 928-936.

Beehler, J., S. Lohr, De­Fo­liart. 1992. Fac­tors in­flu­enc­ing ovipo­si­tion in Aedes tris­e­riatus (Diptera:Culi­ci­dae). The Great Lakes En­to­mol­o­gist, 25/4: 259-264.

CDC, 2009. "La Crosse En­cephali­tis" (On-line). Cen­ters for Dis­ease Con­trol and Pre­ven­tion. Ac­cessed De­cem­ber 06, 2013 at http://​www.​cdc.​gov/​LAC/​index.​html.

Car­pen­ter, S., W. La Casse. 1955. Mos­qui­toes of North Amer­ica. Berke­ley, Cal­i­for­nia: Uni­ver­sity of Cal­i­for­nia Press.

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