Zaprionus tuberculatus

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Zaprionus tuberculatus
Zaprionus (8112935457).jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Family: Drosophilidae
Genus: Zaprionus
Subgenus: Zaprionus
Species:
Z. tuberculatus
Binomial name
Zaprionus tuberculatus
Malloch, 1932

Zaprionus tuberculatus (commonly known as the vinegar fly or the pomace fly) is a member of the subgenus and genus Zaprionus , family Drosophilidae, and order Diptera. [1] It is an invasive fruit fly that originated in Africa, [2] but can also be found in Europe and Asia. [3] The fly earned its common name, the "vinegar fly", because researchers frequently captured the species using vinegar traps. [3] Z. tuberculatus was previously considered a strictly tropical fly, but evidence of invasion to nontropical regions such as Turkey has been shown. [3] [4]

Contents

This fly typically resides on rotting fruit, and the larvae have been found to develop on 49 species of fruits. [3] The life cycle of the fly depends on the temperature of the environment as males are sterile at or above 30 degrees Celsius. It is reddish-brick colored and contains four longitudinal stripes down its head and body. [2] [5] The courtship behavior includes song singing and complex dance rituals. [6] [7] In regard to other organisms, Z. tuberculatus is considered a secondary pest to fruit and a potential threat to ecosystems it invades. [8] A close relative, Z. indianus is an invasive fly species that infests fig orchards.

Taxonomy

Phylogeny of the genus Drosophila and included genera

Sophophora and included groups

Simplified phylogenetic tree of the genus Drosophila and included genera with a focus on the phylogenetic position of the genus Zaprionus.

Zaprionus tuberculatus was first discriminated from its close relative Z. sepsoides in the 1970s. [1] Z. tuberculatus has recently classified as a member of the Tuberculatus species subgroup and is a member of the inermis species group. [3] [5] Previously, Z. tuberculatus was classified as a member of the armatus group. [1] The genus Zaprionus if primarily found throughout Africa and contains 59 species, the most common of which being Z. indianus and Z. tuberculatus. [2] The 59 species of the genus Zaprionus are divided into two subgroups: Afrotropical Zaprionus sensu stricto (s.s.), to which Z. tuberculatus belongs, and Anaprionus near Asia and Australia. [9] [5] Within the Zaprionus sensu stricto (s.s.) subgenus are several further subgroups, including the armatus subgroup to which Z. tuberculatus belonged until 2010. [5]

Description

Zaprionus tuberculatus contains four white horizontal stripes across its head and thorax, similar to other members of the Zaprionus subgroup. The males bear hairs on its forelegs. [2] [5] Both male and female flies have a protruding bristle from the forefemur. [2] The frons have a medium-white stripe and the aedeagus is robust and curved [5] and they have a dark-brown colored thorax. [2] Two white, aligned, horizontal lines characterize the mesonotum and metanotum. [2] Z. tuberculatus shares several common features with other members of the genus Zaprionus, including a completely smooth larval cephalopharyngeal. [5]

Reproductive system

Between individual flies, siblings are distinguished by testicular size (TST), for which the mean is 3.2mm. [2] [5] The average seminal recepaical (SR) of a female fly is 3.6mm. [5]

Distribution

Zaprionus tuberculatus, originally from islands near the Indian Ocean and Afrotropical Region, then became invasive and expanded geographically to southern Europe. [2] Within the Afrotropical region, the species has been reported in Mauritius, Réunion, Madagascar, Saint-Helena, Cape Verde, and Seychelles . [9] The species had first been reported in Europe in Italy September, 2013. [9] Z. tuberculatus has also been reported in North Africa, Malta, Cyprus, and the Canary Islands. [9] Z. tuberculatus was trapped in Eastern Europe in Southern Romania between September and October 2014. [3] Z. tuberculatus has been reported in a few other areas of Europe, including Spain, Greece, and Northern Italy. [3] Z. tuberculatus has also been reported in Asian regions, including Israel and southern Turkey. [3] [4]

Habitat

Vinegar trap similar to the tool used to capture Z. tuberculatus in the wild Fruit fly trap.jpg
Vinegar trap similar to the tool used to capture Z. tuberculatus in the wild

Zaprionus tuberculatus typically reside on rotting fruit and has been captured via the use of vinegar traps, which earned the species its nickname "vinegar fly." [3] The fly does not thrive in strictly tropical environments.

Ecology

Zaprionus tuberculatus is an invasive species, meaning its spread from one geographical region to other may harm to existing ecosystems. [9]

Life cycle

Zaprionus tuberculatus life cycle varies depending on the temperature of the environment. Zaprionus tuberculatus males are sterile at or above 30 degrees Celsius. The mean Z. tuberculatus female life span varies from about 40 days to 60 days, whereas the mean male lifespan varies from about 50 to 200 days when temperature is varied.

Egg

The eggs of Z. tuberculatus have four filaments, with the posterior filaments being elongated more than the anterior filaments and are spatulated. [5] The lengths of these filaments vary within species.

Larvae

The larvae of Z. tuberculatus generally grows on over-ripe fruits and has been found to develop on 49 species of fruits. [9] Similar to other drosophilids, the larvae of Z. tuberculatus have a respiratory system containing thoracic and anterior spiracles on either side of the body, which is a systems organization referred to as amphipneustic. [5]

Puparium

The pupae of Z. tuberculatus have a brick red color. [5] The anterior branches belonging to Z. tuberculatus are clubbed type and vary in number from 11 to 14. [5] The puparial shape (Puparial length:PI) of Z. tuberculatus is 2.59. Puparial shape is an important measurement because it distinguishes Z. tuberculatus from several closely related fly species, including Z. inermis (puparial shape =2.62), Z. cercus (puparial shape =2.40), and Z. burlai (puparial shape = 2.29). [5] Similarly, another taxonomic measurement, the horn index (H) discriminates Z. tuberculatus (H= 7.0) from closely related fly species such as Z. verruca (H=10.6) and Z. burlai (H = 7.2). [5] Unlike other species of Drosophila such as D. melanogaster , for Z. tuberculatus pupae synapsis is uncommon and tend to pupariate upwards relate to the ground instead of downwards. However, Z. tuberculatus pupae strongly aggregate similar to other Drosophila species.

Behavior

Laboratory behavior

Zaprionus tuberculatus pupae are difficult to study experimentally because, when space in culture bottles become limited, pupae climb upwards to escape from the bottle and later die. [5] This is true for all species of the genus Zaprionus, except Zaprionus lachaisei. [5]

Pupation site selection

Though kin selection theory predicts Z. tuberculatus pupae should be more likely to aggregate in an intraspecific manner than interspecifically, Z. tuberculatus does not kin discriminate in site selection.

Species recognition

"Type 2" songs are sung between two male Z. tuberculatus flies and its function is species recognition. [6]

Mating

Male courtship behavior

The male Z. tuberculatus generally makes the first contact with the female Z. tuberculatus by grappling at the female, performing mating displays, then auditory display (song singing). [6] The male performs stationary displays from the female Z. tuberculatus rear end at degrees of 90, 180, or 135 along the long axis of the female body. [6] Still while singing, the male Z. tuberculatus then moves radially around the female in a quarter circle. [6] From a further distance from the female compared to the circling display, the male Z. tuberculatus also performs a dancing which interrupts periods of singing or occurs after singing, but never before. [6] Immediately preceding copulation, then for a second time at the beginning of copulation the male Z. tuberculatus also rubs his legs against the abdomen of the female Z. tuberculatus. [6] The average length of Z. tuberculatus copulation is about 2.25 minutes. [6]

Male songs

Male Z. tuberculatus produces two types of songs by vibrating its wings to generate audio. [7] The two song types are designated either "type 1," which is involved in male courtship, or "type 2," which is used for species recognition both in male and female Z. tuberculatus flies. [7] When one male mounts another, both males sing, possibly to recognize sex or intimidation tactic to deter other males from attempting to mate with the female Z. tuberculatus flies.

Female courtship behavior

At the conclusion of copulation, female Z. tuberculatus protrudes the abdomen and secreted a colorless liquid, which appears to repulse the male Z. tuberculatus. [6] The chemical composition of the colorless liquid has not been studied.

Genetics

The Z. tuberculatus genome has five rod-shaped chromosomes and one dot-shaped chromosome. [10] The number of type of chromosomes, or the karyotype, for Z. tuberculatus has not changed much throughout evolution, which supports Muller's linkage conservation hypothesis and explains the scarcity of pericentric inversions and translocations. [10] [11]

The transposable element mariner occurs in several species, including the Melanogaster species and in the genus Zaprionus. [12] The presence of mariner in the Z. tuberculatus occurs through horizontal transfer. Mariner is also present in several members of the melanogaster species subgroup. The rate of divergence of the mariner element between the two species subgroups suggests a slower evolution rate of the mariner transposable element. [12] A slowing mariner mutation rate helps to explain low divergence in the melanogaster species subgroup, but fails to account for reduced divergence in Z. tuberculatus. The specific mechanism by which horizontal transfer of mariner occurs for Z.tuberculatus is currently unknown and is being investigated. Similarly, study of the retrosposon copia also revealed evidence supporting horizontal transfer between Z. tuberculatus and the melanogaster species subgroup. [13]

Interactions with humans

Zaprionus tuberculatus is a pest in regions to which it has invaded and infects fruit. [8] Z. tuberculatus is thus capable of negatively impacting fruit which humans eat, but it is unknown which fruit species are most negatively influenced. A close relative of Z. tuberculatus, Z. indianus, is an invasive fly species and a pest to fig orchards. However, the adverse ecological influences of Z. tuberculatus has only recently been investigated directly. [14]

Related Research Articles

<i>Drosophila</i> Genus of flies

Drosophila is a genus of fly, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit. They should not be confused with the Tephritidae, a related family, which are also called fruit flies ; tephritids feed primarily on unripe or ripe fruit, with many species being regarded as destructive agricultural pests, especially the Mediterranean fruit fly.

<i>Drosophila melanogaster</i> Species of fruit fly

Drosophila melanogaster is a species of fly in the family Drosophilidae. The species is often referred to as the fruit fly or lesser fruit fly, or less commonly the "vinegar fly", "pomace fly", or "banana fly". In the wild, D. melanogaster are attracted to rotting fruit and fermenting beverages, and are often found in orchards, kitchens and pubs.

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

The Drosophilidae are a diverse, cosmopolitan family of flies, which includes species called fruit flies, although they are more accurately referred to as vinegar or pomace flies. Another distantly related family of flies, Tephritidae, are true fruit flies because they are frugivorous, and include apple maggot flies and many pests. The best known species of the Drosophilidae is Drosophila melanogaster, within the genus Drosophila, also called the "fruit fly." Drosophila melanogaster is used extensively for studies concerning genetics, development, physiology, ecology and behaviour. Many fundamental biological mechanisms were discovered first in D. melanogaster. The fruit fly is mostly composed of post-mitotic cells, has a very short lifespan, and shows gradual aging. As in other species, temperature influences the life history of the animal. Several genes have been identified that can be manipulated to extend the lifespan of these insects. Additionally, Drosophila subobscura, also within the genus Drosophila, has been reputed as a model organism for evolutionary-biological studies, along with D. sechellia for the evolution of host specialization on the toxic noni fruit and Scaptomyza flava for the evolution of herbivory and specialist on toxic mustard leaves.

<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.

<span class="mw-page-title-main">Drosophilinae</span> Subfamily of flies

The Drosophilinae are the largest subfamily in the Drosophilidae. The other subfamily is the Steganinae.

<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.

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.

Fruit fly may refer to:

<i>Drosophila suzukii</i> Species of fly

Drosophila suzukii, commonly called the spotted wing drosophila or SWD, is a fruit fly. D. suzukii, originally from southeast Asia, is becoming a major pest species in America and Europe, because it infests fruit early during the ripening stage, in contrast with other Drosophila species that infest only rotting fruit.

<i>Drosophila</i> (subgenus) Subgenus of flies

Drosophila is a paraphyletic subgenus of the genus Drosophila, a classification of fruit flies. This subgenus was first described by Alfred Sturtevant in 1939. Members of the subgenus Drosophila can be distinguished from other Drosophilid species by breaks in the pigmentation along the dorsal section of their abdomen.

<i>Zaprionus</i> Genus of flies

The genus Zaprionus belongs to the family fruit fly Drosophilidae and is positioned within the paraphyletic genus Drosophila. All species are easily recognized by the white longitudinal stripes across the head and thorax. The genus is subdivided in two subgenera, based on the presence of an even or odd number of white stripes. The species of the genus can be found in Africa and Southern Asia. One species, Zaprionus indianus, has invaded the New World.

<i>Drosophila subobscura</i> Species of insect (fruit fly)

Drosophila subobscura is a species of fruit fly in the family Drosophilidae. Originally found around the Mediterranean, it has spread to most of Europe and the Near East. It has been introduced into the west coasts of Canada, the United States, and Chile. Its closest relative is Drosophila madeirensis, found in the Madeira Islands, followed by D. guanche, found in the Canary Islands. These three species form the D. subobscura species subgroup. When they mate, males and females perform an elaborate courtship dance, in which the female can either turn away to end the mating ritual, or stick out her proboscis in response to the male's, allowing copulation to proceed. D. subobscura has been regarded as a model organism for its use in evolutionary-biological studies.

<i>Scaptomyza flava</i> Species of fly

Scaptomyza flava is an herbivorous leaf mining fly species in the family Drosophilidae. In Latin, flava means golden or yellow. The fly is amber to dark brown in color and approximately 2.5 mm in length. In Europe and New Zealand the larvae are pests of plants in the order Brassicales, including arugula, brassicas, broccoli, Brussels sprouts, bok choy, cabbage, canola, cauliflower, horseradish, kale, kohlrabi, napa cabbage, nasturtium, radish, rapini, rutabaga, turnip, wasabi and watercress. In New Zealand, its range has expanded to include host species that are intercropped with salad brassicas, including gypsophila, otherwise known as baby's breath, which is in the pink family (Caryophyllaceae) and the pea in the Fabaceae. More typically, S. flava is oligophagous within the Brassicales. Scaptomyza are unusual within the Drospophilidae because the group includes species that are truly herbivorous. Other herbivorous drosophilids include D. suzukii, which attacks fruit very early during ripening and species within the genus Lordiphosa, from Africa and Asia, which also include leaf miners. Most drosophilids feed on microbes associated with decaying vegetation and sap fluxes.

Drosophila elegans is a flower-feeding species of fruit flies, belonging to the family Drosophilidae. It is found in Taiwan and the Philippines in Asia.

<i>Zaprionus indianus</i> Species of fly

Zaprionus indianus, the African fig fly, is a species of vinegar fly in the family Drosophilidae.

<i>Drosophila quinaria</i> species group Species group of the subgenus Drosophila

The Drosophila quinaria species group is a speciose lineage of mushroom-feeding flies studied for their specialist ecology, their parasites, population genetics, and the evolution of immune systems. Quinaria species are part of the Drosophila subgenus.

<i>Drosophila silvestris</i> Species of fly

Drosophila silvestris is a large species of fly in the family Drosophilidae that are primarily black with yellow spots. As a rare species of fruit fly endemic to Hawaii, the fly often experiences reproductive isolation. Despite barriers in nature, D. silvestris is able to breed with D. heteroneura to create hybrid flies in the laboratory.

Hirtodrosophila mycetophaga is a fairly large drosophilid fly, with a mean length of 4.0–4.5 mm. It has thus far only been found in Australia. It mates on bracket fungi, preferentially those with a lighter-colored surface in order to enhance mating displays. In addition to these physical displays, flies emit specific sounds in order to attract and ultimately copulate with females.

Hawaiian <i>Drosophila</i> Group of flies

The Hawaiian Drosophilidae are a lineage of flies within the genus Drosophila. This monophyletic clade includes all of the endemic Hawaiian Drosophila and all members of the genus Scaptomyza, which contains both Hawaiian and non-Hawaiian species. The Hawaiian Drosophilidae are descended from a common ancestor estimated to have lived 25 million years ago. Species of Hawaiian Drosophilidae flies have been studied as models of speciation and behavioral evolution. Along with other members of the native Hawaiian ecosystem, the conservations status of many species of Hawaiian Drosophilidae is threatened by habitat loss and introduced predators, among other factors.

Drosophila ochrobasis is an endangered species of fly from Hawaii, in the species rich lineage of Hawaiian Drosophilidae. It is found on the Big island of Hawaii, and has historically been recorded from four of the five volcanoes, though it is now largely absent from most of those sites. This species is in the adiastola subgroup and is closely related to D. setosimentum, but male D. ochrobasis have strikingly different wing markings.

References

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  2. 1 2 3 4 5 6 7 8 9 Patlar, B., Koc, B., Yilmaz, M. and Ozsov, E. D. 2012. First records of Zaprionus tuberculatus (Diptera: Drosophilidae) from the Mediterranean Region, Turkey. Drosophila Information Service 95: 94–96.
  3. 1 2 3 4 5 6 7 8 9 Chireceanu, Constantina; Teodoru, Andrei; Chiriloaie, Andrei (2015-01-01). "The First Detection of Fruit Fly Zaprionus Tuberculatus Malloch (Diptera: Drosophilidae) in the Eastern Part of Europe (Romania)". Annals of the University of Craiova. XX (LVI): 377–382.
  4. 1 2 KUYULU, Akın; YÜCEL, Seda; GENÇ, Hanife (2019-09-02). "Molecular identification of invasive drosophilid Zaprionustuberculatus Malloch (Diptera: Drosophilidae) in Çanakkale, Turkey". Turkish Journal of Zoology. 43 (5): 531–535. doi: 10.3906/zoo-1905-6 . ISSN   1303-6114.
  5. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Yassin, Amir; David, Jean R. (2010-07-23). "Revision of the afrotropical species of Zaprionus (Diptera, Drosophilidae), with descriptions of two new species and notes on internal reproductive structures and immature stages". ZooKeys (51): 33–72. doi: 10.3897/zookeys.51.380 . ISSN   1313-2989. PMC   3088026 . PMID   21594121.
  6. 1 2 3 4 5 6 7 8 9 Bennet-Clark, H. C.; Leroy, Y.; Tsacas, L. (1980-02-01). "Species and sex-specific songs and courtship behaviour in the genus Zaprionus (Diptera-Drosophilidae)". Animal Behaviour. 28 (1): 230–255. doi:10.1016/S0003-3472(80)80027-3. ISSN   0003-3472.
  7. 1 2 3 Lee, R. C. P. (1986). "The role of male song in sex recognition in Zaprionus tuberculatus (Diptera, Drosophilidae) - Washington University in St Louis". Animal Behaviour. 34 (3): 641–648. doi:10.1016/S0003-3472(86)80050-1 . Retrieved 2019-10-31.
  8. 1 2 "Zaprionus tuberculatus". https. Retrieved 2019-10-31.
  9. 1 2 3 4 5 6 Raspi, Alfio; Grassi, Alberto; Benelli, Giovanni (2014-06-01). "Zaprionus tuberculatus (Diptera Drosophilidae): First records from the European mainland". Bulletin of Insectology. 61.
  10. 1 2 Su, Y.; Herrick, K.; Farmer, J. L.; Jeffery, D. E. (1992-07-01). "Zaprionus tuberculatus: Chromosome Map and Gene Mapping by DNA in situ Hybridization". Journal of Heredity. 83 (4): 299–304. doi:10.1093/oxfordjournals.jhered.a111216. ISSN   0022-1503. PMID   1401876.
  11. ↑ Muller HJ, 1940. Bearings of the Drosophila work on systematics. In: The new systematics (Huxley J, ed). Oxford: Clarendon Press: 185-268.
  12. 1 2 Maruyama, Kyoko."Interspecific Transfer of the Transposable Element Mariner Between Drosophila and Zaprionus," (PhD diss., Washington University in St.Louis School of Medicine,1991).
  13. ↑ De Almeida, Luciane M; Carareto, Claudia MA (2006). "Sequence heterogeneity and phylogenetic relationships between the copia retrotransposon in Drosophila species of the repleta and melanogaster groups". Genetics Selection Evolution. 38 (5): 535–50. doi: 10.1186/1297-9686-38-5-535 . ISSN   1297-9686. PMC   2689262 . PMID   16954045.
  14. ↑ Leivadaras J, Leivadara EI, Vontas J (2017) A new insect, drosophila Zaprionus tuberculatus (Diptera: Drosophilidae), affects the figs on Crete. Abstract of a paper presented at the Panhellenic Entomological Congress (Athens, GR, 2017-09-19/22), p 27.
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