Anastrepha ludens

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Anastrepha ludens
Anastrepha ludens 1322089.jpg
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 OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Family: Tephritidae
Genus: Anastrepha
Species:
A. ludens
Binomial name
Anastrepha ludens
Loew, 1873
Synonyms
  • Acrotoxa ludens
  • Anastrepha lathana
  • Trypeta ludens

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]

Contents

Anastrepha ludens is native to Mexico and Central America and is a major pest to citrus and mango agriculture in Mexico, Central America, and the lower Rio Grande Valley. [2] The species exhibits high fecundity and relatively long lifespans compared to other species of fruit flies. [3] These qualities make the Mexican fruit fly a particularly aggressive invasive species, especially threatening agriculture because the larvae grow and feed on many different species of fruit. [4] The Anastrepha genus is designated as one of three genera that pose the greatest risk to American agriculture. [5] According to the USDA, A. ludens is the only important member of the Anastrepha genus that is subtropical instead of tropical and thus has a range much further North than most Anastrepha species. [2] Also the USDA estimates that the Mexfly causes $1.44 billion worth of damage in a 5-year time span, mostly to citrus farms. [6] They are frequently designated as an invasive species in Southern California and Arizona and pose a serious threat to Florida's grapefruit agriculture. [4]

Description

Adults

Left: Female A. ludens. Right: male A. ludens. Anastrepha ludens female and male.jpg
Left: Female A. ludens. Right: male A. ludens.

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 mesonotum that is 2.75-3.6 mm long and a wing span of 6.6-9.0 mm. [2]

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]

Anastrepha ludens pupae Anastrepha-ludens-pupae.webp
Anastrepha ludens pupae

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]

Anastrepha ludens larva.jpg
Larva of Anastrepha ludens
Anastrepha ludens larva anal lobes.jpg
Anterior spiracle of a larval Anastrepha ludens
Anastrepha ludens larva posterior spiracle.jpg
Anastrepha ludens larva posterior spiracle
Anastrepha ludens larva anterior spiracle.jpg
Anastrepha ludens larva anterior spiracle
Anastrepha-ludens-pupae.webp
Anastrepha ludens pupa

Distribution

Anastrepha ludens is native to Guatemala, Mexico and possibly Costa Rica. [2] It is an invasive species to the US. [4] The first record of these flies spotted outside of their native habitat of Mexico and Central America was in a small Texas colony in 1903. By 1927, Mexican fruit flies were infecting citrus farms in lower Rio Grande Valley of Southern Texas and threatening farmland in California, Texas, Florida, and Arizona. In 1954, the fly species had spread westward as far as Hermosillo. [11] The flies were rare in Costa Rica until the 1990s when they suddenly appeared on citrus plants. It is unclear if the species was native to Costa Rica or migrated there from the north. It was previously believed that the species is native to Colombia because of misidentification of A. manizaliensis but it is now known that the species does not exist there. [12]

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]

Oranges Ambersweet oranges.jpg
Oranges

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:

[15] [16]




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

Lateral view of female (left) and male (right) Anastrepha ludens. Anastrepha ludens female and male.jpg
Lateral view of female (left) and male (right) Anastrepha ludens.

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. [3] After this period the male is fully sexually active. The female fly can lay over 1500 eggs in its lifetime. [4] These flies are known to be able to go through period of estivation. This ability and their polyphagous nature allows them to be able to survive in poor resource conditions better than other flies as they migrate to find a site with better resources. [3] Anastrepha ludens female reproductive potential has been shown to be affected by male-female contact. An experiment showed that combining females and males together in cages during maturation reduced egg production. [17]

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 olfactory and visual stimulus to find a good oviposition site. They have been observed landing on potential host fruit and walking around while headbutting the fruit. Once the female makes her decision to lay eggs, she will bore into the fruit and deposit eggs. Then she deposits a host-marking pheromone over her eggs. This pheromone seems to stimulate the female fly.

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]

Related Research Articles

<span class="mw-page-title-main">Sterile insect technique</span> Method of biological control for insect populations

The sterile insect technique (SIT) is a method of biological insect control, whereby overwhelming numbers of sterile insects are released into the wild. The released insects are preferably male, as this is more cost-effective and the females may in some situations cause damage by laying eggs in the crop, or, in the case of mosquitoes, taking blood from humans. The sterile males compete with fertile males to mate with the females. Females that mate with a sterile male produce no offspring, thus reducing the next generation's population. Sterile insects are not self-replicating and, therefore, cannot become established in the environment. Repeated release of sterile males over low population densities can further reduce and in cases of isolation eliminate pest populations, although cost-effective control with dense target populations is subjected to population suppression prior to the release of the sterile males.

<span class="mw-page-title-main">Apple maggot</span> Species of fly

The apple maggot, also known as the railroad worm, is a species of fruit fly, and a pest of several types of fruits, especially apples. This species evolved about 150 years ago through a sympatric shift from the native host hawthorn to the domesticated apple species Malus domestica in the northeastern United States. This fly is believed to have been accidentally spread to the western United States from the endemic eastern United States region through contaminated apples at multiple points throughout the 20th century. The apple maggot uses Batesian mimicry as a method of defense, with coloration resembling that of the forelegs and pedipalps of a jumping spider.

<span class="mw-page-title-main">Tephritidae</span> Family of fruit flies

The Tephritidae are one of two fly families referred to as fruit flies, the other family being the Drosophilidae. The family Tephritidae does not include the biological model organisms of the genus Drosophila, which is often called the "common fruit fly". Nearly 5,000 described species of tephritid fruit fly are categorized in almost 500 genera of the Tephritidae. Description, recategorization, and genetic analyses are constantly changing the taxonomy of this family. To distinguish them from the Drosophilidae, the Tephritidae are sometimes called peacock flies, in reference to their elaborate and colorful markings. The name comes from the Greek τεφρος, tephros, meaning "ash grey". They are found in all the biogeographic realms.

<i>Bactrocera tryoni</i> Species of fly

Bactrocera tryoni, the Queensland fruit fly, is a species of fly in the family Tephritidae in the insect order Diptera. B. tryoni is native to subtropical coastal Queensland and northern New South Wales. They are active during the day, but mate at night. B. tryoni lay their eggs in fruit. The larvae then hatch and proceed to consume the fruit, causing the fruit to decay and drop prematurely. B. tryoni are responsible for an estimated $28.5 million a year in damage to Australian crops and are the most costly horticultural pest in Australia. Up to 100% of exposed fruit can be destroyed due to an infestation of this fly species. Previously, pesticides were used to eliminate B. tryoni from damaging crops. However, these chemicals are now banned. Thus, experts devoted to B. tryoni control have transitioned to studying this pests' behaviors to determine a new method of elimination.

<i>Ceratitis capitata</i> Species of insect

Ceratitis capitata, commonly known as the Mediterranean fruit fly or medfly, is a yellow-and-brown fly native to sub-Saharan Africa. It has no near relatives in the Western Hemisphere and is considered to be one of the most destructive fruit pests in the world. There have been occasional medfly infestations in California, Florida, and Texas that require extensive eradication efforts to prevent the fly from establishing itself in the United States.

<i>Bactrocera dorsalis</i> Species of insect

Bactrocera dorsalis, previously known as Dacus dorsalis and commonly referred to as the oriental fruit fly, is a species of tephritid fruit fly that is endemic to Southeast Asia. It is one of the major pest species in the genus Bactrocera with a broad host range of cultivated and wild fruits. Male B. dorsalis respond strongly to methyl eugenol, which is used to monitor and estimate populations, as well as to annihilate males as a form of pest control. They are also important pollinators and visitors of wild orchids, Bulbophyllum cheiri and Bulbophyllum vinaceum in Southeast Asia, which lure the flies using methyl eugenol.

<i>Rhagoletis mendax</i> Species of fly

Rhagoletis mendax is a species of tephritid fruit fly known by the common name blueberry maggot. The blueberry maggot is closely related to the apple maggot, a larger fruit fly in the same genus. It is a major pest of plant species in the Ericaceae family, such as blueberry, cranberry, and huckleberry. The larva is 5 to 8 mm long, apodous, and white with chewing mouthparts. Female adults are 4.75 mm in length, males are slightly smaller. Both adults are mostly black in color with white stripes, orange-red eyes, and a single pair of clear wings with black banding. The adult female fly lays a single egg per blueberry, and when the larva hatches it consumes the fruit, usually finishing the entire berry in under 3 weeks and rendering it unmarketable. The larva then falls to the soil and pupates. Adult flies emerge, mate, and females oviposit when blueberry plants are producing fruit. Each female fly can lay 25 to 100 eggs in their lifetime.

<i>Rhagoletis cerasi</i> Species of fly

Rhagoletis cerasi is a species of fruit fly in the family Tephritidae.

<i>Bactrocera cucurbitae</i> Species of fly

Bactrocera cucurbitae, the melon fly, is a fruit fly of the family Tephritidae. It is a serious agricultural pest, particularly in Hawaii.

<i>Anastrepha</i> Genus of flies

Anastrepha is the most diverse genus in the American tropics and subtropics. Currently, it comprises more than 300 described species, including nine major pest species, such as the Mexican fruit fly, the South American fruit fly, the West Indian fruit fly, the sapote fruit fly, the Caribbean fruit fly, the American guava fruit fly, and the pumpkin fruit fly, as well as the papaya fruit fly. As some of their names suggest, these pest species are one of the most numerous and damaging groups of insects in their native range, plaguing commercial fruits such as citrus, mango, guava, and papaya.

Rhagoletis juglandis, also known as the walnut husk fly, is a species of tephritid or fruit fly in the family Tephritidae. It is closely related to the walnut husk maggot Rhagoletis suavis. This species of fly belongs to the R. suavis group, which has a natural history consistent with allopatric speciation. The flies belonging to this group are morphologically distinguishable.

<span class="mw-page-title-main">False codling moth</span> Species of moth in family Tortricidae

Thaumatotibia (Cryptophlebia) leucotreta, commonly known as the false codling moth, orange moth, citrus codling moth or orange codling moth, is a moth in the family Tortricidae under the order of Lepidoptera. Larvae of the moth feed on a wide range of crops from cotton and macadamia nuts to Citrus species. The larvae have a less selective diet than the codling moth, which feeds primarily on temperate fruit crops.

<i>Planococcus citri</i> Species of true bug

Planococcus citri, commonly known as the citrus mealybug, is a species of mealybugs native to Asia. It has been introduced to the rest of the world, including Europe, the Americas, and Oceania, as an agricultural pest. It is associated with citrus, but it attacks a wide range of crop plants, ornamental plants, and wild flora.

<i>Anastrepha suspensa</i> Species of fly

Anastrepha suspensa, known as the Caribbean fruit fly, the Greater Antillean fruit fly, guava fruit fly, or the Caribfly, is a species of tephritid fruit fly. As the names suggest, these flies feed on and develop in a variety of fruits, primarily in the Caribbean. They mainly infest mature to overripe fruits. While thought to have originated in Cuba, the Caribbean fruit fly can now also be found in Florida, Hispaniola, and Puerto Rico.

Ceratitis rosa, or commonly known as the natal fruit fly, is a fruit fly species from the family Tephritidae of the order Diptera. It is considered a pest in both its native and non-native habitats. It prefers subtropical or temperate climates and the fruits that are native to these areas. It is also capable of surviving in less dry climates compared to its relative, the Ceratitis capitata which is more specific in its habitats, described by Anton Ferdinand Karsch in 1887 in Delagoa Bay, Mozambique. Not long after in 1900, the natal fruit fly was a recognized pest for orchards and fruit farmers throughout KwaZulu Natal Province, Republic of South Africa.

<i>Diachasmimorpha longicaudata</i> Species of wasp

Diachasmimorpha longicaudata is a solitary species of parasitoid wasp and an endoparasitoid of tephritid fruit fly larvae. D. longicaudata is native to many countries in Southeast Asia and subtropical regions and has also been introduced to many other countries as a biological control agent. It is now considered the most extensively used parasitoid for biocontrol of fruit flies in both the southern portion of the United States and Latin America. D. longicaudata is especially useful for agricultural purposes in the control of fruit flies as it is easily mass-reared and has the ability to infect a variety of hosts within the genus Bactrocera. A negative factor in its use as a biocontrol agent is that it is known to oviposit in grapefruit in the state of Florida. This has resulted in quarantines on grapefruit shipped internationally as well as domestically. Research is ongoing to determine whether D. longicaudata is actually a single species, or if it contains multiple species. It is likely multiple biological species separated by both reproductive isolation and morphological characteristics such as wing geometry.

<i>Paracantha gentilis</i> Species of fly

Paracantha gentilis is a species of tephritid or fruit fly in the genus Paracantha of the family Tephritidae. It has a widespread distribution throughout the Western United States, and has also been found as far south as Mexico and Costa Rica. It most closely resembles Paracantha culta, which is widespread in the Southeastern United States, but P. gentilis can be distinguished by having smaller spots on the head.

<i>Bactrocera carambolae</i> Species of fly

Bactrocera carambolae, also known as the carambola fruit fly, is a fruit fly species in the family Tephritidae, and is native to Asia. This species was described by Drew and Hancock in 1994.

Bactrocera passiflorae, the Fijian fruit fly, is a species of fly in the family Tephritidae in the insect order Diptera. It is native to several tropical and subtropical islands in the Pacific Ocean and is a pest of fruit crops.

<i>Anastrepha fraterculus</i> South American fruit fly

Anastrepha fraterculus, known as the South American fruit fly, is a fruit fly species from the genus Anastrepha. A. fraterculus is a polyphagous, frugivorous fly that is a significant pest of commercial fruit production in South America.

References

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Sources

Identification