''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
Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
to
Southeast Asia
Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
. It is one of the major pest species in the genus ''Bactrocera'' with a broad
host range
In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasite, parasitic, a mutualism (biology), mutualistic, or a commensalism, commensalist ''guest'' (symbiont). The guest is typically provided with ...
of cultivated and wild fruits. Male ''B. dorsalis'' respond strongly to
methyl eugenol
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. ...
, 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.
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''.
Description
''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
The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen.
In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
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
The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
.
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
The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...
for depositing eggs in host fruits, while in male adults this
ovipositor
The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...
is notably absent.
Distribution
Endemic to
Southeast Asia
Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
, ''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
Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
, the
Mariana Islands
The Mariana Islands ( ; ), also simply the Marianas, are a crescent-shaped archipelago comprising the summits of fifteen longitudinally oriented, mostly dormant volcanic mountains in the northwestern Pacific Ocean, between the 12th and 21st pa ...
, and
Tahiti
Tahiti (; Tahitian language, Tahitian , ; ) is the largest island of the Windward Islands (Society Islands), Windward group of the Society Islands in French Polynesia, an overseas collectivity of France. It is located in the central part of t ...
. The fly is also found in most countries of sub-Saharan Africa. 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.
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. In July 2010, flies were discovered in traps in the
Sacramento
Sacramento ( or ; ; ) is the capital city of the U.S. state of California and the seat of Sacramento County. Located at the confluence of the Sacramento and American Rivers in Northern California's Sacramento Valley, Sacramento's 2020 p ...
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.
CLIMEX, a modeling software, has been used to map the future trajectory for the fruit fly in terms of opportunities for increasing its distribution. This was tested both under current and future predicted climate conditions, given the current research on
climate change
Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
. 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. 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.
Habitat
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.
They can be found in the
USA
The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 states and a federal capital district, Washington, D.C. The 48 contiguous ...
,
China
China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
, the
UAE
The United Arab Emirates (UAE), or simply the Emirates, is a country in West Asia, in the Middle East, at the eastern end of the Arabian Peninsula. It is a federal elective monarchy made up of seven emirates, with Abu Dhabi serving as i ...
, the rest of
Southeast Asia
Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
,
India
India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
,
Brunei
Brunei, officially Brunei Darussalam, is a country in Southeast Asia, situated on the northern coast of the island of Borneo. Apart from its coastline on the South China Sea, it is completely surrounded by the Malaysian state of Sarawak, with ...
,
Malaysia
Malaysia is a country in Southeast Asia. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...
, the
Philippines
The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
, The
Mariana Islands
The Mariana Islands ( ; ), also simply the Marianas, are a crescent-shaped archipelago comprising the summits of fifteen longitudinally oriented, mostly dormant volcanic mountains in the northwestern Pacific Ocean, between the 12th and 21st pa ...
,
Timor Leste
Timor-Leste, also known as East Timor, officially the Democratic Republic of Timor-Leste, is a country in Southeast Asia. It comprises the eastern half of the island of Timor, the coastal exclave of Oecusse in the island's northwest, and ...
,
Pakistan
Pakistan, officially the Islamic Republic of Pakistan, is a country in South Asia. It is the List of countries and dependencies by population, fifth-most populous country, with a population of over 241.5 million, having the Islam by country# ...
,
Taiwan
Taiwan, officially the Republic of China (ROC), is a country in East Asia. The main geography of Taiwan, island of Taiwan, also known as ''Formosa'', lies between the East China Sea, East and South China Seas in the northwestern Pacific Ocea ...
and the rest of
South Asia
South Asia is the southern Subregion#Asia, subregion of Asia that is defined in both geographical and Ethnicity, ethnic-Culture, cultural terms. South Asia, with a population of 2.04 billion, contains a quarter (25%) of the world's populatio ...
. One male fly was recently found in
New Zealand
New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
, but it is so far unclear as to whether a breeding population has been established.
Life history
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. 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. Flies typically live from 1–3 months, but this can be extended to up to 12 months under cool conditions.
Eggs
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.
Larvae
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.
Pupae
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. Pupation can take approximately 10–12 days, but depending on the season, this process can be delayed in cooler conditions.
Adults
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.
Food resources
''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.
Social behavior
Mating
Polyandry
Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives ...
has been observed in ''B. dorsalis''. 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. 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.
Reproductive
senescence
Senescence () or biological aging is the gradual deterioration of Function (biology), functional characteristics in living organisms. Whole organism senescence involves an increase in mortality rate, death rates or a decrease in fecundity with ...
does appear to be present in this species, as male and female age correlates negatively with the rate of fertilization.
Flying
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.
Transmission electron microscopy
Transmission electron microscopy (TEM) is a microscopy technique in which a beam of electrons is transmitted through a specimen to form an image. The specimen is most often an ultrathin section less than 100 nm thick or a suspension on a g ...
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.
Long-distance flight
''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.
Symbiotic relationships
''B. dorsalis'' has
symbiotic relationships
Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term
with many bacteria. Different bacteria dominate at different developmental stages of the fruit fly.
Pseudomonadota
Pseudomonadota (synonym "Proteobacteria") is a major phylum of gram-negative bacteria. Currently, they are considered the predominant phylum within the domain of bacteria. They are naturally found as pathogenic and free-living (non- parasitic) ...
are most often present in immature stages, whereas ''
Bacillota
The Bacillota (synonym Firmicutes) are a phylum of bacteria, most of which have Gram-positive cell wall structure. They have round cells, called cocci (singular coccus), or rod-like forms (bacillus). A few Bacillota, such as '' Megasphaera'', ...
'' are most often present in the adult stages. Overall, the most abundant families are ''
Enterococcaceae
The Enterococcaceae are a family of Gram-positive bacteria placed in the order Lactobacillales.
Representative genera include ''Enterococcus'', ''Melissococcus'', ''Pilibacter'', ''Tetragenococcus'', and ''Vagococcus''. In this family are some ...
Comamonas
''Comamonas'' is a genus of bacteria in the phylum Pseudomonadota.Garrity, George M.; Brenner, Don J.; Krieg, Noel R.; Staley, James T. (eds.) (2005). Bergey's Manual of Systematic Bacteriology, Volume Two: The Proteobacteria, Part C: The Alpha- ...
'' are extremely abundant in pupae, but disappear entirely by adulthood.
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.
Aposymbiotic
Aposymbiosis occurs when symbiotic organisms live apart from one another (for example, a clownfish living independently of a sea anemone). Studies have shown that the lifecycles of both the host and the symbiont are affected in some way, usually ...
flies responded faster to diets in experimental conditions and fed more, for longer periods of time.
Interactions with humans
Human activities are mainly 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. 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.
Agricultural pest
''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.
Conventional pest treatment
''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. 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.
To resolve these concerns, several techniques have been implemented, including
sterile insect technique
The sterile insect technique (SIT) is a method of biological pest control, biological insect control, whereby overwhelming numbers of infertility, sterile insects are released into the wild. The released insects are preferably male, as this is mo ...
, protein bait sprays, and male annihilation. 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.
One of the most experimentally effective control techniques has been the wrapping of fruit, often in a paper or
polythene
Polyethylene or polythene (abbreviated PE; IUPAC name polyethene or poly(methylene)) is the most commonly produced plastic. It is a polymer, primarily used for packaging (plastic bags, plastic films, geomembranes and containers including bott ...
sleeve. This physical barrier prevents oviposition from occurring. 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.
Parasitoid wasps
In addition to these, Hawaii has developed methods to suppress ''Bactrocera'' species using
parasitoid wasp
Parasitoid wasps are a large group of hymenopteran Superfamily (zoology), superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, ...
s, 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.
Field sanitation
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.
Insecticide resistance
Many organophosphorus insecticides target the enzyme
acetylcholinesterase
Acetylcholinesterase (HUGO Gene Nomenclature Committee, HGNC symbol ACHE; EC 3.1.1.7; systematic name acetylcholine acetylhydrolase), also known as AChE, AChase or acetylhydrolase, is the primary cholinesterase in the body. It is an enzyme th ...
(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''.
References
Further reading
* Allwood, A.J., Chinajariyawong, A., Drew, R.A.I., et al. (1999) Host plant records for fruit flies (Diptera: Tephritidae) in south east Asia. ''Raffles Bulletin of Zoology Supplement 7:1-92.
United States National Agricultural Library
The United States National Agricultural Library (NAL) is one of the world's largest agricultural research libraries, and serves as a national library of the United States and as the library of the United States Department of Agriculture. Locate ...