Anopheles quadrimaculatus
(common malaria mosquito)

Pictures

Picture	Title	Caption	Copyright
Title Adult Caption Anopheles quadrimaculatus (common malaria mosquito); adult at rest. At one time, the most important vector of malaria in the eastern United States and, recently, was found to be a host for the West Nile Virus in the USA. Copyright Public Domain - Released by Centers for Disease Control & Prevention (CDC) - Original image by Edward McCellan, 1962.	Adult	Anopheles quadrimaculatus (common malaria mosquito); adult at rest. At one time, the most important vector of malaria in the eastern United States and, recently, was found to be a host for the West Nile Virus in the USA.	Public Domain - Released by Centers for Disease Control & Prevention (CDC) - Original image by Edward McCellan, 1962.
Title Breeding grounds Caption Anopheles quadrimaculatus (common malaria mosquito); water hyacinth in a Louisiana (USA) pond can act as a breeding ground for Anopheles quadrimaculatus, Mansonia and Culex mosquitoes, which anchor onto the plants, and are protected from Copyright Public Domain - Released by Centers for Disease Control & Prevention (CDC) - Original image by Graham Heid & Dr. Harry D. Pratt	Breeding grounds	Anopheles quadrimaculatus (common malaria mosquito); water hyacinth in a Louisiana (USA) pond can act as a breeding ground for Anopheles quadrimaculatus, Mansonia and Culex mosquitoes, which anchor onto the plants, and are protected from	Public Domain - Released by Centers for Disease Control & Prevention (CDC) - Original image by Graham Heid & Dr. Harry D. Pratt

Identity

Preferred Scientific Name

• Anopheles quadrimaculatus Say

Preferred Common Name

• common malaria mosquito

Other Scientific Names

• Anopheles annulimanus Wulp, 1867

Local Common Names

• Germany: Gabelmücke

Summary of Invasiveness

A. quadrimaculatus a mosquito is the chief vector of malaria in North America. This species prefers habitats with well-developed beds of submergent, floating leaf or emergent aquatic vegetation. Larvae are typically found in sites with abundant rooted aquatic vegetation, such as rice fields and adjacent irrigation ditches, freshwater marshes and the vegetated margins of lakes, ponds and reservoirs. 

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Arthropoda
•             Subphylum: Uniramia
•                 Class: Insecta
•                     Order: Diptera
•                         Family: Culicidae
•                             Genus: Anopheles
•                                 Species: Anopheles quadrimaculatus

Description

Anopheles quadrimaculatus is described as a large, dark brown mosquito. The tarsus is entirely dark (The Ohio State University Mosquito Pest Management Bulletin,1998). O'Malley (1992) reports that, "All Anopheles adults are characterized by an evenly rounded scutellum and palpi about as long as the proboscis. A. quadrimaculatus is a medium-sized species. Wings are entirely dark scaled and 4 mm or more in length. Scutal bristles are short and wings are spotted with patches of dark scales. The tip of the wing is dark without copper-colored fringe scales. The palpi have dark scales and are unbanded, and the wing has 4 distinct dark-scaled spots .."

Distribution

Native range: North America; Anopheles quadrimaculatus has a distribution that covers much of the eastern United States. Its range extends from southern Canada to the Florida Everglades, and to the west from Minnesota to Mexico (Kaiser, 1994). Please follow this link for a distribution map (Levine et al. 2004).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Habitat

Chase and Knight (2003) state that, "Many species of mosquitoes are habitat generalists which breed, grow as larvae and emerge from a wide variety of aquatic habitats .." O'Malley (1992) reports that, "In North America, most anophelines prefer habitats with well-developed beds of submergent, floating leaf or emergent aquatic vegetation. Larvae are typically found in sites with abundant rooted aquatic vegetation, such as rice fields and adjacent irrigation ditches, freshwater marshes and the vegetated margins of lakes, ponds and reservoirs. Investigators have suggested that aquatic vegetation promotes anopheline production because it provides a refuge for larvae from predators, such as Gambusia affinis. Additional hypotheses for the beneficial effects of aquatic vegetation include: enhanced food resources in vegetated regions, shelter from physical disturbance and favorable conditions for oviposition (Orr and Resh 1989)."

Habitat List

Host Animals

Species Vectored

Biology and Ecology

Nutrition
O'Malley (1992) reports that, "A. quadrimaculatus larvae are indiscriminate feeders whose natural food includes a wide range of aquatic organisms, both plant and animal, as well as detritus. This food may be living or dead at the time of ingestion. The main criterion in selecting food seems to be whether the suspended material is small enough to eat. When feeding, A. quadrimaculatus larvae lie horizontally, with the dorsal side just under the surface film. The head rotates 180 degrees horizontally so that it is actually upside down and the venter of the head is dorsal. Feeding is either "eddy feeding" or "interfacial feeding". Eddy feeding is employed for infusions when the surface contains islets of floating oil materials. Two eddies with converging streams unite in front of the larva to form a current toward the mouth from a distance of about half the length of the larva. Efferent currents flow outward at right angles to the body from the antenna. Particles too large to eat are held by the maxillae, drawn below the surface and discarded as the head is rotated to the normal position. Interfacial feeding on the membranes of algae, bacteria, debris and fungi is common in nature. Feeding in this manner is accomplished by setting up currents which draw particles to the mouth from all directions in a straight line and at nearly equal velocities. Surface tension of the larval habitat determines the type of feeding. Eddy feeding occurs at a surface tension of less than 60 dynes per square cm; interfacial feeding is practiced in habitats with a surface tension above 62 dynes per square cm." O'Malley (1992) reports that, "Mosquito feeding patterns are largely regulated by host availability and preference (Apperson and Lanzaro 1991). Female A. quadrimaculatus are primarily mammalian feeders and actively feed on man and on wild and domesticated animals. As noted previously, this is a significant pest species. Females repeatedly seek their hosts, often visiting the same feeding site several times during the course of a bloodmeal."

Chase and Knight (2003) state that, "Larvae of the two most common mosquito species encountered in the natural and artificial wetlands, A. quadrimaculatus and C. pipiens, and other types of mosquito larvae, utilize different feeding behaviours and have slightly different diets (e.g. Merritt et al. 1992). They are both generalists, however, and readily consume detritus, microbes and algae, both from the benthos and the water column. As such, they are likely to compete for resources with several other co-occurring species."

Reproduction
The Ohio State University Mosquito Pest Management Bulletin (1998) reports that, "Anopheles quadrimaculatus eggs are laid singly on the water surface with lateral floats to keep them at the surface. One hundred or more eggs are laid at a time. A single female may lay as many as 12 batches of eggs and a total of more than 3,000 eggs." O'Malley (1992) reports that, "Mating occurs as soon as the females emerge. Males wait in nearby vegetation and seek females as they begin to fly. Copulation is completed in flight and takes 10-15 seconds. One insemination is usually sufficient for the fertilization of all eggs."

Lifecycle stages
Floore (2004) states that, "The mosquito goes through four separate and distinct stages of its life cycle: egg, larva, pupa, and adult. Each of these stages can be easily recognized by its special appearance." Egg stage: Eggs are laid one at a time or attached together to form "rafts." They float on the surface of the water. In the case of Culex and Culiseta species, the eggs are stuck together in rafts of up to 200. Anopheles, Ochlerotatus and Aedes , as well as many other genera, do not make egg rafts, but lay their eggs singly. Culex, Culiseta, and Anopheles lay their eggs on the water surface while many Aedes and Ochlerotatus lay their eggs on damp soil that will be flooded by water. Most eggs hatch into larvae within 48 hours; others might withstand subzero winters before hatching. Water is a necessary part of their habitat.

Larval stage: The larva (plural - larvae) lives in the water and comes to the surface to breathe. Larvae shed (molt) their skins four times, growing larger after each molt. Most larvae have siphon tubes for breathing and hang upside down from the water surface. Anopheles larvae do not have a siphon and lie parallel to the water surface to get a supply of oxygen through a breathing opening. Coquillettidia and Mansonia larvae attach to plants to obtain their air supply. The larvae feed on microorganisms and organic matter in the water. During the fourth molt the larva changes into a pupa (Floore, 2004).

Pupal stage: The pupal stage is a resting, non-feeding stage of development, but pupae are mobile, responding to light changes and moving (tumble) with a flip of their tails towards the bottom or protective areas. This is the time the mosquito changes into an adult. This process is similar to the metamorphosis seen in butterflies when the butterfly develops - while in the cocoon stage - from a caterpillar into an adult butterfly. In Culex species in the southern United States this takes about two days in the summer. When development is complete, the pupal skin splits and the adult mosquito (imago) emerges (Floore, 2004).

Adult:: The newly emerged adult rests on the surface of the water for a short time to allow itself to dry and all its body parts to harden. The wings have to spread out and dry properly before it can fly. Blood feeding and mating does not occur for a couple of days after the adults emerge (Floore, 2004).

Impact Summary

Impact

A. quadrimaculatus can also transmit Cache Valley virus (CV) (Blackmore et al), West Nile Virus (CDC, 2007) and transmission of St. Louis encephalitis has been obtained with this species in laboratory experiment (Horsfall 1972 in O’Malley, 1992).

A. quadrimaculatus has been found to be an excellent host for dog heartworm (Dirofilaria immitis). According to Lewandowski et al. (1980), this is probably one of the most important species involved in the natural transmission of dog heartworm in Michigan. In central New York, this species was also the most efficient host of dog heartworm out of several species tested, both in the laboratory and the wild (Todaro and Morris 1975).

A. quadrimaculatus can be a vector for the myositic parasite Trachipleistophora hominis. Weidner et al. (1999) found that, Microsporidian spores of T. hominis Hollister, isolated from a human, readily infected larval stages of both A. quadrimaculatus. The authors state that, "Nearly 50% of the infected mosquito larvae survived to the adult stage. Spores recovered from adult mosquitoes were inoculated into mice and resulted in significant muscle infection at the site of injection".

Risk and Impact Factors

• Invasive in its native range
• Proved invasive outside its native range
• Benefits from human association (i.e. it is a human commensal)
• Has high reproductive potential

• Host damage
• Negatively impacts human health
• Negatively impacts animal health
• Negatively impacts tourism

• Causes allergic responses
• Pest and disease transmission

Similarities to Other Species/Conditions

Levine et al. (2004) report that, "A. quadrimaculatus was considered to be a single species until biological evidence necessitated subdividsion into a species complex in the late 1900s. A combination of genetic crossing, isozyme, and ctytological information convincingly showed that there are at least five species in the group and they include: A. quadrimaculatus, A. smaragdinus, A. diluvialis, A. inundatus, and A. maverlius." The A. quadrimaculatus complex as a whole is often referred to as A. quadrimaculatus (sensu lato), whereas A. quadrimaculatus (sensu stricto) refers to the individual species (Rios and Connelly, 2008). The authors state that A. quadrimaculatus is the most widely distributed of the species complex in the eastern United States and southeastern Canada (Seawright et al. 1991)."

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Chemical Control

A study by Dennett et al (2003) compared the effects of fipronil and lambda-cyhalothrin on A. quadrimaculatus larvae in Arkansas rice plots, USA. Results showed that at 24 hours post-treatment, higher control was obtained for A. quadrimaculatus with fipronil, and was less harmful to non-target predators.

Ham et al. (1999) compared mortality of Anopheles quadrimaculatus, Culex quinquefasciatus and Aedes spp. complex after exposure to the insecticides Dibrom, Trumpet and Scourge in Louisiana, USA. “At all time intervals, Dibrom and Trumpet were significantly more effective against the Aedes spp. complex than against An. Quadrimaculatus and Cx. quinquefasciatus. Scourge was significantly more effective against An. Quadrimaculatus and Cx. quinquefasciatus than Dibrom or Trumpet” (Ham et al. 1999).

Biological Control

Effectiveness of Bacillus larvicides were also compared in Arkansas rice plots during the 1998 growing season. Experimental Bacillus larvicides designed to float on or near the water surface were compared to labeled standard Bacillus corn-cob-based larvicides. “Experimental floating formulations of Bacillus thuringiensis var israelensis applied at 5.58 and 11.18 kg/ha provided up to 100% control of 3rd- and 4th-stage Anopheles larvae within 24-48 h, whereas the water-dispersible granule formulations containing Bacillus sphaericus required 48-72 h to yield >75% mortality in 0.16-ha plots at 11.18 kg/ha.” (Dennett and Meisch 2000).

Cyclopoid copepods are currently in use for control against Aedes larvae in Louisiana, USA, and thus may offer possibilities for Anopheles control. Rice fields in Louisiana are a major breeding habitat for Anopheles quadrimaculatus. Marten et al (2000) measured how natural and introduced copepod populations affect Anopheles larvae in the rice fields. Naturally occurring copepod predators present in almost all fields were Mesocyclops ruttneri, Acanthocyclops vernalis and Macrocyclops albidus. In fields where these copepods were present A. quadrimaculatus larvae production was suppressed due to predation. Macrocyclops albidus, M. ruttneri, Mesocyclops edax and Mesocyclops longisetus were introduced to experimental rice field plots. “It took two months for the introduced copepods to build up their numbers; Anopheles larvae then disappeared from all treated plots while larvae continued to be present in the adjacent control field.” The authors conclude that it seems likely that Anopheles production can be reduced by using cultivation practices that encourage predator populations and introducing select species of copepods (Marten et al. 2000).
 

Bibliography

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References

ISSG, 2011. Global Invasive Species Database (GISD). Invasive Species Specialist Group of the IUCN Species Survival Commission. http://www.issg.org/database

Distribution References

CABI Data Mining, 2001. CAB Abstracts Data Mining.,

CABI, Undated. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

Invasive Species Specialist Group (ISSG), 2011. Global Invasive Species Database (GISD). In: Global Invasive Species Database (GISD), Auckland, New Zealand: University of Auckland. http://www.issg.org/database

Principal Source

Global Invasive Species Database

Contributors

Principal sources:Shiff, 2002 Integrated Approach to Malaria Control
Levine et al. 2003. Distribution of Members of Anopheles quadrimaculatus Say s.l. (Diptera: Culicidae) and Implications for Their Roles in Malaria Transmission in the United States.

Distribution Maps