Anastrepha ludens
Anastrepha ludens | |
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Anastrepha ludens on their preferred orange fruit habitat. The bottom right fly is a female as identified by long ovipositor. The fly in the middle of image is male. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Diptera |
Family: | Tephritidae |
Genus: | Anastrepha |
Species: | A. ludens
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Binomial name | |
Anastrepha ludens Loew, 1873
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Synonyms | |
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Anastrepha ludens, the Mexican fruit fly or Mexfly,[1] is a species of fly of the Anastrepha genus in the Tephritidae family (fruit flies). It is closely related to the Caribbean fruit fly Anastrepha suspensa, and the papaya fruit fly Anastrepha curvicauda.[2]
Anastrepha ludens is native to
Description
Adults
Most species in the Anastrepha genus including A. ludens have a distinctive yellow and brown coloration of the body and wings. The adult A. ludens is 7–11 mm long, or slightly larger than a common house fly. They have a
Females have a relatively long life spans of up to 11 months. Males live even longer than females, up to 16 months.[7] Female adult A. ludens have a long ovipositor (3.35-4.7mm) and sheath relative to body size and are capable of laying more than 1,500 eggs in their lifetimes,[7] making A. ludens highly fecund.
Anastrepha ludens have 12 chromosomes and most cells are diploid.[8] A 2014 genetic study of A. ludens concluded that "A. ludens populations are genetically diverse with moderate levels of differentiation." They go on to attribute this genetic diversity to natural selection across the wide habitat range of the fly and to pest management practices. In addition they found high levels of inbreeding in the species.[9]
Gut bacteria exist in A. ludens called Enterobacteriaceae (fruit fly type bacteria). Other families of bacteria have been found in Mexican fruit flies including Vibrionaceae, Bacillaceae, Micrococcaceae, and Pseudomonadaceae.[10] The effects of these bacteria on A. ludens are not well studied but it has been proposed by M. Aluja that A. ludens regurgitate internal bacteria onto their host and use the bacterial colonies as a protein source. The gut bacteria may also play a role in digestion and detoxification of chemicals.[3]
Larvae
Larvae have an elongated cylindrical shape typical of fruit flies and are white in color. Larvae can be up to 12 mm in length.[2] Larvae usually pupate on the ground but have also been observed to occasionally pupate inside its host fruit. Larvae determine when to exit a fruit through physical and chemical signals such as the pH of the rotting fruit and the drop of the fruit from to the ground.[3]
Distribution
Anastrepha ludens is native to
The US Department of Agriculture presumes that these flies were introduced to the USA through the import of larvae infested fruits.[5]
Currently, California, Arizona, Florida and most of northern Mexico are considered free of Mexican fruit flies and eradication efforts are taking place in Texas,[13][14] with some Texas quarantine areas having been successfully cleared and the quarantines lifted.[1] The eradication of these flies from most of the US and Northern Mexico is largely due to the successful implementation of the sterile insect technique (SIT). SIT is currently used in parts of Texas to control the species population.[13]
Habitat
The Mexican fruit fly prefers living near citrus and other fruits, which act as hosts where the female can lay her eggs. Anastrepha suspensa prey on many of the same fruits in the same regions where A. ludens primarily reside as well. Anastrepha ludens prefers laying eggs in grapefruits and oranges, but many other hosts have been recorded including:
- Anacardium occidentale(cashew)
- Annona cherimola(cherimoya)
- Annona reticulata (custard apple)
- Annona squamosa (sugar-apple)
- Carica papaya(papaya)
- Casimiroa edulis(white sapote)
- Casimiroa tetrameria (matasano)
- Citrus aurantiifolia(lime)
- Citrus maxima(pummelo)
- Citrus aurantium(sour orange)
- Citrus limetta (sweet lime)
- Citrus x paradisi(grapefruit)
- Citrus medica(citron)
- Citrus reticulata(tangerine)
- Citrus sinensis(sweet orange)
- Coffea arabica (arabica coffee)
- Cydonia oblonga(quince)
- Diospyros kaki (Japanese persimmon)
- Feijoa sellowiana (feijoa)
- Inga (shimbillo)
- Malus domestica(apple)
- Malus pumila(paradise apple)
- Mammea americana (mammey apple)
- Mangifera indica (mango)
- Sideroxylon capiri (bully tree)
- Passiflora edulis (purple granadilla)
- Persea americana(avocado)
- Pouteria sapota (sapote)
- Prunus persica(peach)
- Psidium guajava (common guava)
- Psidium littorale(strawberry guava)
- Punica granatum(pomegranate)
- Pyrus communis (pear)
- Sargentia greggii (yellow chapote)
- Spondias purpurea (red mombin)
- Syzygium jambos (rose-apple)
It has been experimentally shown that A. ludens choose oranges and grapefruit over other hosts but in the absence of these fruits will deposit larvae on any of the above hosts.[4]
Life history
While female and male A. ludens can live up to 11 and 16 months respectively under lab conditions, in nature their lifespans tend to be about a year.[3] Females typically lay approximately 25-70 eggs a day. Their relatively long life span allows females to have a gross reproduction rate of up to 1600 offspring.[3] They prefer to lay their eggs on citrus fruits, typically grapefruits or oranges, when the fruits start to ripen and develop in color.[7]
The Mexican fruit fly goes through four stages of development (holometabolism or complete metamorphosis): egg, larvae, pupa, adult. The rate at which they mature is directly related to ambient environmental factors such as temperature and humidity.[11]
The life cycle begins when the adult female lays her eggs. The female fly deposits eggs via her ovipositor into the fruit host. The eggs hatch 6–10 days later and then enter their second stage of development, the larval stage.[3]
The larval stage lasts for 3–4 weeks, depending on the temperature and other conditions. While inside the fruit, the larvae continue to grow and develop through 3 larval instars. When they have reached optimal size and environmental conditions are right, the mature larvae emerge from the fruit into the soil and begin to pupate.[3] Larvae eat and burrow into the fruit that their mother laid them on. While grapefruits and oranges are preferred, other citrus fruits, pears, apples, and peach are also common hosts and thus food sources.[4] Like other fruit flies, A. ludens need to consume a mixture of amino acids, minerals, carbohydrates, water, and vitamins in order to survive. Adult flies can get their nutrients from raindrops and bird feces, while larvae get all their resources from their host fruit.[3]
Adult
The adult fly emerges from the pupal casing and the life cycle begins anew. The new fly finds a dry sheltered spot until it can unfold their wings. There is a period of sexual maturation during which they eat lots of protein which allows for gonadal development.
Mating and parental care
Male A. ludens exhibit lek mating and thus do not provide any care for offspring outside of fertilizing the egg.[3]
Female A. ludens will use
Anastrepha ludens males follow a lek mating strategy in which they provide no parental care for offspring. Males mating strategy involves claiming a territory and defending it from other males through sounds and physical actions. Ideal territories for males are under the leaves of trees that produce citrus fruit. Males deposit their pheromones through their mouth and anus onto the underside of leaves, and they emit an aggressive song by quickly vibrating their wings.[3]
Female A. ludens exhibit mate choice and tend to prefer to mate with larger males. This is thought to be due to larger males singing better mating songs and depositing more sperm into females. The process of a female choosing a mate can take up to 2 hours.[3]
Social behavior
Larvae will feed on their host fruit for continuous periods of over 24 hours. Larval movement is dictated by the ripeness of the host fruit. After the larvae matures to become an adult, 96% of A. ludens emerge from their burrow hole between 6 a.m. and 10 a.m. Mating calls of adult flies are observed mostly during late afternoons. These behaviors vary between different species of fruit flies.[18]
A. ludens have been observed migrating about 135 km from their breeding site in Mexico to farms in southern Texas. Using release-recapture technique, researchers observed flies moving back and forth between the two habitat areas.[19]
Ecology
Predators
The main natural enemies of A. ludens are parasitoid wasps, specifically in the families Braconidae and Ichneumonidae.[20] Diachasmimorpha longicaudata, Doryctobracon crawfordi, Ganaspis pelleranoi, Biosteres giffardi, B. vandenboschi, and Aceratoneuromyia indica have been released by the governments of the US, Costa Rica, Mexico, Brazil, Argentina[failed verification] and Peru to biologically control A. ludens and other Anastrepha species populations.[21] D. longicaudata and D. crawfordi have been established as the most efficient at controlling population size for this species of flies.[3]
Interaction with humans
According to the USDA the species is "one of the world's most destructive invasive pests". In 2018, the Texas citrus industry supported nearly 6,000 jobs and accounted for more than $465 million in statewide revenue. [22]
The USDA utilizes integrated pest management tactics to control the threat of an invasion. The USDA implements a quarantine zone where wild flies are captured in the United States. Movement of citrus fruit is restricted within the quarantined area. The USDA operates an extensive eradication and suppression campaign against the Mexican fruit fly (Mexfly), which includes the use of sterile insect technique. Sterile flies are released by the hundreds of millions to suppress the invasive population.[23] Other tactics deployed by the USDA include the use of preventative pesticide application and biocontrol tactics by releasing parasitoid wasps, which are natural A. ludens predators.[5] There has been at least one Mexfly quarantine in Texas on an annual basis for over 80 years.[6] The Texas Department of Agracalture (TDA) advises anyone with citrus trees to pick fruits before they fall to the ground to help prevent the spread of the species.[6]
As of July 2021, there is an active A. ludens quarantine zones in the Texas.[24]
References
- ^ a b "Anastrepha ludens Mexican Fruit Fly APHIS Removes the Quarantine Area in Laredo Webb County and Zapata Zapata County Texas". North American Plant Protection Organization's Phytosanitary Alert System. 2020-09-13. Retrieved 2020-12-21.
- ^ a b c d e f Stone, Alan (1942). "The Fruitflies of the Genus Anastrepha". USDA Miscellaneous Publications (439).
- ^ .
- ^ a b c d e f Weens Jr., H.V.; Heppner, J.B.; Gary J., Steck. "Mexican fruit fly (Anastrepha ludens)". Featured Creatures Entomology & Nematology Department. University of Florida. Retrieved 18 September 2019.
- ^ a b c United States Department of Agriculture, Animal and Plant Health Inspection Service Fruit Fly Exclusion and Detection Program (May 2019). "Fruit Fly Exclusion and Detection Strategic Plan FY 2019-2023" (PDF): 1–18.
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: Cite journal requires|journal=
(help) - ^ a b c "The Mexican Fruit Fly". Citrus Alert. Retrieved 2019-12-04.
- ^ PMID 16154309.
- PMID 25472896.
- ISSN 0015-4040.
- S2CID 19952298.
- ^ a b Flitters, N.E. (1965). Effect of temperature and humidity on development and potential distribution of the Mexican fruit fly in the United States. Technical bulletin / United States Department of Agriculture ;no. 1330. Washington: U.S. Dept. of Agriculture.
- ^ "Anastrepha ludens (Mexican fruit fly)". Invasive Species Compendium. CABI.org. Retrieved 31 October 2019.
- ^ a b "Anastrepha ludens (Mexican fruit fly)". Invasive Species Compendium. CABI.org. Retrieved 31 October 2019.
- ^ "Fruit Flies Quarantine Information". United States Department of Agriculture Animal and Plant Health Inspection Service. Retrieved 2020-12-21.
- .
- ^ White, IM; Elson-Harris, MM (1994). "Fruit Flies of Economic Significance: Their Identification and Bionomics". CABI. CAB International: 601.
- PMID 14994799.
- ^ Beal, J.A. (July 1932). "Control of the turpentine border in the stores region". Circular. 226: 20–25.
- ISBN 9780849312755.
- ^ Leyva, Jorge L.; Harold W., Browning; Gilstrap, Frank E. "Effect of Host Fruit Species, Size, and Color on Parasitization of Anastrepha ludens (Diptera: Tephritidae) by Diachasmimorpha longicaudata (Hymenoptera: Braconidae)". Department of Entomology, Texas A&M University.
- ISBN 0851994342. Retrieved 31 October 2019.
- ^ "USDA APHIS | USDA Asks for the Public's Help to Protect Citrus in Texas".
- PMID 17598535. Retrieved 31 October 2019.
- ^ "Fruit Flies Quarantine Information". aphis.usda.gov. Retrieved 31 October 2019.
- Sources
- US Department of Agriculture, APHIS Fact Sheet, Mexican Fruit Fly, Doc. (1993).
External links
- Species Profile- Mexican Fruit Fly (A. ludens). National Invasive Species Information Center United States National Agricultural Library. Lists general information and resources for Mexican Fruit Fly.
- Five-year strategy plan 2008-2013 for fruit flies of Mexico.
- "Fruit Flies Quarantine Information". United States Department of Agriculture Animal and Plant Health Inspection Service. Retrieved 2020-12-21.
Identification
- The Diptera site Comprehensive guide to identification literature.