Bactrocera dorsalis | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Diptera |
Family: | Tephritidae |
Genus: | Bactrocera |
Species: | B. dorsalis |
Binomial name | |
Bactrocera dorsalis (Hendel, 1912) | |
Bactrocera dorsalis range (by country) | |
Synonyms | |
Dacus dorsalis |
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. [1] 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. [2] [3] [4] 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. [5] [6]
The fly is similar to the closely related species B. carambolae and B. occipitalis. The species name B. dorsalis is identical to other synonyms B. papayae, B. invadens and B. philippinensis. [7] [8] [9]
B. dorsalis is a species of tephritid fruit fly. Flies that belong to this family are usually small to medium-sized with colorful markings. In particular, B. dorsalis belongs to a complex of physically similar flies called the Bactrocera dorsalis complex, whose defining characteristics include a mostly black thorax and dark T-shaped marking on the fly's abdominal segment. The T-shape marking consists of a dark medial and transverse band along the fly's abdomen. [10]
The B. dorsalis species has distinctive yellow and black markings on its thorax and abdomen, which may vary between flies. Two vertical yellow markings on the thorax and the dark T-shaped marking on the abdomen differentiate this species of fly from its close relatives. The wings are clear with a continuous costal band. The adult body is around 8.0 mm in length, with wings approximately 7.3 mm in length. The female adult has a tapered ovipositor for depositing eggs in host fruits, while in male adults this ovipositor is notably absent. [11]
Endemic to Southeast Asia, B. dorsalis is a highly invasive pest species that now has a presence in at least 65 countries. It has also been introduced to Hawaii, the Mariana Islands, and Tahiti. The fly is also found in most countries of sub-Saharan Africa. [11] From 1910 to 1990, the fly species was only observed in 5 countries; however, in the last three decades, the rate of spread by B. dorsalis has sharply increased, with the species invading an additional 70 countries. [12]
Elsewhere in the United States, B. dorsalis has been spotted in California and Florida. These appearances then trigger a cascade of eradication efforts. Four major oriental fruit fly eradication efforts occurred in response to infestations in California between 1960 and 1997. Two additional infestations were eradicated in 2006 and 2007, occurring within 3–4 years of reports of these infestations. [11] In July 2010, flies were discovered in traps in the Sacramento and Placer counties of California. A quarantine was established, and eradication efforts followed. These sightings in the mainland United States are generally quarantined infestations that have been eradicated. [13]
CLIMEX, a modeling software, has been used to map the future trajectory for the fruit fly in terms of opportunities for increasing its distribution. [14] This was tested both under current and future predicted climate conditions, given the current research on climate change. Under current conditions, the fly's projected distribution includes much of the tropics and subtropics and extends into areas like Mediterranean Europe. The model predicts optimal climate conditions in the southeast United States. [14] Under climate change conditions, the spread overall increases as the fly is less limited by cold weather. However, its distribution does possibly decrease in areas where precipitation decreases. [14]
These tephritid fruit flies are found in tropical areas. B. dorsalis also prefers to pupate in shaded rather than brightly lit areas, moist over dry soil, and in soil with larger particles (particle size greater than or equal to 2.5 mm) than in soil with smaller particles. [15]
Under summer conditions, development from egg to adult requires 16 days. Several stages can be delayed in cooler conditions. B. dorsalis eggs may take up to 20 days to hatch under cool conditions, extended from the usual single day. [13] Pupariation occurs in the soil under the host plant and is normally completed within 10–12 days, but can be delayed up to 90 days under cool conditions. [13] Flies typically live from 1–3 months, but this can be extended to up to 12 months under cool conditions. [13]
Mated female B. dorsalis puncture the skin of mature fruit and deposit eggs in a few batches of 3-30 eggs underneath a fruit's skin via ovipositor, depending on the quality and ripeness of the fruit. Eggs usually hatch within a day, although the hatching process can be delayed in cool conditions. Females are capable of laying over 3,000 eggs during their lifetime under optimal conditions, but in field conditions, reports a typical range from about 1,200 to 1,500 eggs laid per female. [11]
Eggs hatch to larvae and moult twice (there are three larval instars) while feeding on the flesh of the fruit for about 6–35 days. Larvae are creamy white in color with a maggot-like appearance and are about 10 mm in length. [16]
The third instar larvae exit the fruit and burrow into the soil under the host plant to pupate. Larvae generally pupate in the upper 4 cm of the soil. [15] Pupation can take approximately 10–12 days, but depending on the season, this process can be delayed in cooler conditions. [16]
In 1-2 weeks, the adult emerges from the pupae and matures. Most adults emerge from the soil between 7:00 AM and 10:00 AM. Once sexual maturity is reached (which takes approximately 9 days), adults engage in the mating process and the life cycle repeats. The adult lifespan for B. dorsalis is about 90 days, and the flies have been reported to travel up to 30 miles in search of new egg laying sites and food, such as decaying fruit and plant nectar. Adult females prefer to lay eggs in old egg deposit sites on fruits. [11] [13] [16] [17]
B. dorsalis has been seen in more than 200 kinds of fruit and nut plants, but the species lay eggs in mango, papaya, and avocado fruits most often. Adult flies feed on decaying fruit, plant nectar, and other substances during their lifetime and prefer to feed in the morning. [16] [18]
Polyandry has been observed in B. dorsalis. [19] For females, there is typically a re-mating refractory period. The length of this period does not vary based on whether the female is mating with a virgin or non-virgin male. However, when there was a refractory period, females lay more eggs. [19] Females who were exposed to two males continuously without a refractory period in between lay fewer eggs, but still lay more eggs than females with only one male. Therefore, there appears to be a reproductive benefit for females with polyandry. [19]
Reproductive senescence does appear to be present in this species, as male and female age correlates negatively with the rate of fertilization. [19]
The flight capacity of B. dorsalis adult females at various life stages has been observed in order to better understand and prevent their spread. Past research has shown that the species can spread extensively following fruit harvests, with a dispersal radius up to 37 km observed in Hawaii. [20] Transmission electron microscopy was employed to view changes in flight muscle ultrastructures. Researchers observed that flight speed and distance changed with the age of the fruit fly, reaching its maximum capacity at 15 days of age. [20]
B. dorsalis has been observed to be capable of long-distance flight. One study investigated the relationship between flight muscle structure and the flies' flight capacity: as the number of mitochondria increased, myofibril diameter increased, and sarcomere length decreased, the researchers found the fly's flight capacity to be maximized. This particular muscle structure was evident in 15-day-old female adult B. dorsalis flies. [20]
B. dorsalis has symbiotic relationships with many bacteria. Different bacteria dominate at different developmental stages of the fruit fly. [21] Pseudomonadota are most often present in immature stages, whereas Bacillota are most often present in the adult stages. [21] Overall, the most abundant families are Enterococcaceae and Comamondaceae. Meanwhile, Comamonas are extremely abundant in pupae, but disappear entirely by adulthood. [21]
Researchers have also tested the relationship between certain gut symbiotic bacteria and fly foraging behavior and nutrient ingestion. Suppression of the fly's microbiome resulted in changes in the foraging behavior in both male and female flies. [22] Aposymbiotic flies responded faster to diets in experimental conditions and fed more, for longer periods of time. [22]
Human activities are partially responsible for causing the spread of the species from one region to another. The primary risk comes from import of fruit that may contain larvae, either in passenger cargo, or through the smuggling of fruit in passenger baggage or mail. [13] In New Zealand, researchers recorded 7-33 interceptions of fruit flies per year in cargo. Researchers also recorded an additional 10-28 interceptions in passenger baggage. [13]
B. dorsalis is not only a highly invasive species, accidentally introduced to Hawaii from Taiwan during the 1940s in World War II, but also very destructive to crop yield for farmers of various fruits, vegetables, and nuts across the world. The larval stage of the life cycle is the most damaging to fruits because of larval feeding on the soft flesh of fruits. After ovipositing occurs by a female fly, the larvae develop under the skin of the fruit or soft tissues of the plant and begin to feed on the fruit or plant's flesh. Once feeding occurs, other microorganisms can invade the site of larval feeding and cause the fruit to decay faster. Although ripe fruits are believed to be preferred for ovipositing, unripe fruits have served as hosts for eggs as well. [18]
B. dorsalis has created many agricultural issues for humans, especially in the areas where it is endemic. In the Pacific Islands, the fly has restricted the development of a diverse tropical fruit and vegetable industry, necessitated that commercial fruits go through quarantine treatment before shipment, and provided an avenue for their introduction to countries not previously exposed to the fly species. [23] The Miami-Dade County in Florida had to perform the quarantine technique when the insect was detected in fruits there in August 2015; they were eliminated from the area in February 2016. [24]
To resolve these concerns, several techniques have been implemented, including sterile insect technique, protein bait sprays, and male annihilation. [23] Male annihilation technique is effective because methyl eugenol attracts male flies prior to the beginning of their sexual maturation, to an extent of 40 to 50 percent of the flies. [4]
One of the most experimentally effective control techniques has been the wrapping of fruit, often in a paper or polythene sleeve. This physical barrier prevents oviposition from occurring. [13] The caveat with this method is that it must be implemented far in advance of the fruit fly's presence. Alternatively, fruits can be harvested earlier in the season than the flies anticipate; this has proven effective with the mango fruit. [13]
In addition to these, Hawaii has developed methods to suppress Bactrocera species using parasitoid wasps, including Fopius arisanus. The parasitic wasp oviposits its own eggs into B. dorsalis eggs, the parasitoids are reared in the host, and the developed parasitoids emerge in the pupal stage. F. arisanus has been observed to be the most successful example of parasitoid control of B. dorsalis, and researchers are looking to introduce its model of suppression from Hawaii to other areas of the world that the fly affects. [23]
One simple, but effective technique called field sanitation, through which all unmarketable fruits get removed from the fields and disposed as soon as they are observed, so that re-infestation does not occur. [18]
Many organophosphorus insecticides target the enzyme acetylcholinesterase (AChE); mutations in the acetylcholinesterase gene of B. dorsalis have been found to be associated to resistance to such insecticides. Researchers have identified three point mutations in B. dorsalis' gene encoding AChE that generate nonsynonymous changes in the produced amino acid sequence. Two of the point mutations are identical in site to mutations identified in other Bactrocera species, but one of the mutations is specific to B. dorsalis. Widespread use of such insecticides could result in rapid resistance acquisition in populations of B. dorsalis. [25]
Flies are insects of the order Diptera, the name being derived from the Greek δι- di- "two", and πτερόν pteron "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced mechanosensory organs known as halteres, which act as high-speed sensors of rotational movement and allow dipterans to perform advanced aerobatics. Diptera is a large order containing an estimated 1,000,000 species including horse-flies, crane flies, hoverflies, mosquitoes and others, although only about 125,000 species have been described.
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.
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.
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.
A semiochemical, from the Greek σημεῖον (semeion), meaning "signal", is a chemical substance or mixture released by an organism that affects the behaviors of other individuals. Semiochemical communication can be divided into two broad classes: communication between individuals of the same species (intraspecific) or communication between different species (interspecific).
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.
Bactrocera is a large genus of tephritid fruit flies, with close to 500 species currently described and accepted.
Rhagoletis cerasi is a species of fruit fly in the family Tephritidae.
Bactrocera cucurbitae, the melon fly, is a fruit fly of the family Tephritidae. It is a serious agricultural pest, particularly in Hawaii.
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.
Anastrepha ludens, the Mexican fruit fly or Mexfly, is a species of fly of the Anastrepha genus in the Tephritidae family. It is closely related to the Caribbean fruit fly Anastrepha suspensa, and the papaya fruit fly Anastrepha curvicauda.
Methyl eugenol (allylveratrol) is a natural chemical compound classified as a phenylpropene, a type of phenylpropanoid. It is the methyl ether of eugenol and is important to insect behavior and pollination. It is found in various essential oils.
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Bactrocera (Bactrocera) invadens is the name given to tephritid fruit flies that were introduced to East Africa from Sri Lanka and subsequently invaded practically the whole of Sub-Saharan Africa, hence the species name "invadens". It was first shown to be the same biological species as B. dorsalis s.s. by possessing identical sex pheromonal components after consumption of methyl eugenol, and also based on CO1 and rDNA sequences. Subsequently, it was agreed that B. invadens, B. papayae and B. philippinensis be synonymized as B. dorsalis. To counteract its detrimental effects to the fruit business, the industry resorts to cold treatment in order to get rid of the larvae.
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.
An attractant is any chemical that attracts an organism, e.g. i) synthetic lures; ii) aggregation and sex pheromones ; and iii) synomone
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 discovered 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.
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.