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== Life history and cycle ==
== Life history and cycle ==
[[File:Posterior spiracles of Anastrepha suspensa.tif|thumb|Posterior spiracles of ''A. suspensa.''|353.98895263671875x353.98895263671875px]]
[[File:Posterior spiracles of Anastrepha suspensa.tif|thumb|Posterior spiracles, hairs, trunks, openings|alt=]]


In ''A. suspensa,'' as with other fruit flies, there are 4 main stages: [[egg]], [[larva]] (with 3 [[instar]]s or substages), [[pupa]], and adult.
In ''A. suspensa,'' as with other fruit flies, there are 4 main stages: [[egg]], [[larva]] (with 3 [[instar]]s or substages), [[pupa]], and adult.

Revision as of 20:46, 18 November 2019

Anastrepha suspensa
Scientific classification
Kingdom:
Phylum:
Class:
Order:
Family:
Genus:
Species:
A. suspensa
Binomial name
Anastrepha suspensa
(Loew, 1862)

The Caribbean fruit fly (Anastrepha suspensa), also sometimes called the Greater Antillean fruit fly, guava fruit fly, or the Caribfly, is a fly species in the order Diptera.[2] As the name suggests, these flies are often seen hovering around and infecting an array of fruits primarily from the Caribbean. It is common to see them only infest mature to overripe fruits. While thought to originate in Cuba, the Caribbean fruit fly can now also be found in Florida, Hispaniola, and Puerto Rico.[3][4]

The Caribbean fruit fly is a small-sized, brown and yellow fly that is oftentimes distinguished from similar species by the placement of the veins in the wings.[5][6][7] These small parasites reside primarily in tropic areas, which provide sweet fruits for the flies to use as resources for rapid development.[2] While they are small in size, A. suspensa flies continue to negatively impact agricultural output and economic growth . Due to their ability to travel great distances (over 120 km) and wide range of adaptability, the Caribfly poses grave risks to any country importing fruit from the Caribbean.[8]

Description

This adult fly is between 11-14 mm long and may appear yellow, orange, brown, or a mixture of all three.[2] A. suspensa, though very similar to other species in the genus Anastrepha, may be distinguished by the amulets on its ovipositor.[7] On females, the ovipositor is sharp with serrations near the tip.

The Caribbean fruit fly appears primarily yellow and brown with an orange-brown body and a yellow head. The setae, or hairlike structures on insects, are red-brown to dark-brown, causing a darker overall color.[7] Similar to other species in Anastrepha, their thorax is primarily yellow to orange-brown with slender bands of color running down the thorax to the scutum.[7] In addition, dark spots may be found on the thorax and wings of the fly.[2]

Distribution and territory

The Caribbean fruit fly, as the name suggests, has primarily been identified in Caribbean Islands and Central America in countries like the Bahamas, British Virgin Islands, Cuba, Dominican Republic, Haiti, Jamaica, and Puerto Rico. While primarily widespread in these countries, A. suspensa has also been found in California and Florida, as well as in French Guiana in South America.[2][4][9]

A. suspensa have the ability to fly large distances, but will often claim the territory of a leaf in the late afternoon as a means of attracting and mating with females. Here, larger males have an advantage when fighting for territories with the largest males having a greater likelihood of obtaining a larger territory.[10][11]

Habitat

Tropical fruits which A. suspensa. consumes.

Since A. suspensa primarily infests fruits like guava and grapefruit, and they typically live and lay eggs in areas that possess an abundance of these tropical fruits. Therefore, they often reside in forests with suitable fruits.[12] The spread of A. suspensa throughout the globe has largely been attributed to the movement of spoiled fruit that contained A. suspensa larvae.[9][12]

A. suspensa prefer to reside in tropical climates with consistent precipitation. While tropical climates may include anything greater than 1500 mm of precipitation annually to less than 600 mm, there has been no distinct preference between these subcategories by A. suspensa.[citation needed]

Life history and cycle

Posterior spiracles, hairs, trunks, openings

In A. suspensa, as with other fruit flies, there are 4 main stages: egg, larva (with 3 instars or substages), pupa, and adult.

Egg

Caribbean fruit fly eggs are elliptical in shape.[2][4][13] The color of these eggs may range from pale to gleaming white. The anterior end of the egg contains a micropyle, which is an opening for the sperm to enter. Additionally, on this end of the egg, there is a characteristic polygonal pattern in the surface architecture of the chorion. Female Caribbean fruit flies will lay about 200 of their eggs on the exterior of mature and rotting fruit.[14]

Larva

Since the larvae are vermiform maggots, the Caribbean fruit fly larvae mimic many of the other morphological components of other fruit flies due to their wedge shape. Therefore, they possess an elongated and cylindrical shape. These larvae progress through 3 instars, or a stage between two stages of molting in the development. First instar larvae appear clear to pale white. During this period, the mouth hooks are formed and become the main feature.[14] These two sets of thin brown hooks are located on the ventral surface of the larvae. The color of second instar larvae begins to change towards a pale cream color.[14] In this stage, the larvae will begin to develop features like the anterior spiracles to allow oxygen to enter the respiratory system.[14] They will also start to develop small oral ridges and black mouth hooks. Third instar larvae transition from a cream to a light yellow color and are about 8 to 10 mm in length. During this the time, the overall body surface becomes rough and the morphology of the larvae solidifies. The mouth hooks have now transformed into a single pair of sharp, black teeth that are strongly sclerotized.[14] The oral ridges become more distinct, and there are creeping welts that begin to form.[14] Due to the close resemblance of A. suspensa and other species in Anastrepha, it can be difficult the accurately identify A. suspensa larvae.[citation needed]

Puparium

The pupae are fully encompassed by the hardened conveying and skin of the 3rd larval install.[14] The pupae possess an ellipsoid shape and appear golden to reddish-brown. During this phase, there are visible spiracles on the anterior surface that allow for respiration.[citation needed]

Adult

The adult A. suspensa appear yellow and brown and typically range between 12–14 mm in length.[14] Their wings often alternate between bands of yellow-brown and brown bands.[10] Distinguishing A. suspensa adults from other adults in Anastrepha is difficult, but can be done by examining the positioning of the veins on the wings of the flies.[2] On the other hand, male and female A. suspensa can easily be distinguished. Female A. suspensa have a prominent ovipositor, which is the organ through which a female insect deposits her eggs. In A. suspensa, even compared to other species of Anastrepha the ovipositor will be short and thin with serrations at the bottom two-thirds of the tip.[citation needed]

Food resources

A. suspensa on fruit.

A. suspensa have been recorded to infest and consume over 100 foods, most of which have been fruits.[citation needed] These fruits are almost always mature, overripe, or spoiled. These flies seem to prefer consumption of guavas, the Cayenne cherry, rose apple, tropical almonds, grapefruit, and oranges. While these fruits are preferred, they have also been noted to infest papaya, Natal plum, kumquat, mango, mombin, and an array of cherries.[15] The data shows that the consumption of different fruit may depend on the location of the A. suspensa.[citation needed]

Food for A. suspensa plays a role in movement and growth because they are correlated with the maturity of the fruit.[16] Larvae and pupae advance towards the center and pulp sections of the fruit as it matures because of the increased sweetness.[16] This then leads to more rapid development.[16]

Mating

By studying behavioral ecology of A. suspensa, clear mating patterns have been noticed between males and females.

While these studies were conducted during spring and summer, it has been noted that A. suspensa do not have a winter diapause, which is a period of suspended development during the winter, so this mating behavior may be generalizable to the entire year.[17] During the early morning, it is common for males and females to be interacting on fruit like guava and Surinam cherries. The males will be feeding and courting, and the females will be feeding and ovipositing.[17] As temperature and light increase as the morning progresses, males and females will both move to areas under shaded leaves.[17] As the morning turns into afternoon and late afternoon, there will male competition for single-leaf territories. Since territorial fights were a crucial component to mating, flies that were larger and could claim larger territories were able to demonstrate higher levels of fitness and had better matting success.[17] In these claimed areas, the males will puff to produce sex pheromones from glands near their anal canal and sing songs to attract females.[18] In almost all of the situations and samples considered in the experiments, mating occurred after these displays of fitness.[citation needed]

Marking

Marking has also been observed by male A. suspensa. This marking is implemented through scent.[19] Male A. suspensa will perch and then release air-borne pheromones from the tip of their abdomen to mark and defend individual leaves.[19][20]

Songs

It has been noted that male A. suspensa make two distinct sounds towards females: the calling and precopulatory song.[18][21] Both of these sounds are created rapid wing vibrations by the male A. suspensa.[18][21] The various studies have observed these songs to understand differences between different males who produce the sounds. It has been shown that larger size is correlated with more intensity for both songs.[18][21] Male size seem to be negatively correlated with frequency of calling songs, but there is no correlation between male size and frequency of precopulatory songs.[18][21] Additionally, it has been shown that females are attracted to areas where the calling song is broadcast, but more research must be done on whether calling males have higher mating success than males that do not call.[18][21]

Since unsuccessful precopulatory songs were not as loud and had a broader bandwidth than that of successful songs, it is believed that the precopulatory song may be the last acoustic opportunity for males to display their fitness to the female.[18][21] If this prediction is true, it would explain greater breeding for better precopulatory songs due to female compliance once a male has mounted. Interestingly, a longer precopulatory song has been demonstrated to be correlated with shorter matings. The main hypothesis for this occurrence is that longer precopulatory songs are typically for mates that are less sexually responsive; therefore, the longer song may be just enough to lead to mating.[18][21] Since larger males coupled longer, it is believed that the increased strength that comes with size helps in a conflict of interest when it comes to mating between males and females.[18][21]

Parental care

A. suspensa females on average laid about 200 eggs.[10] Studies examining oviposition looked at the influence of oviposition length noted that A. suspensa females experiencing 72 hour oviposition periods mature faster than females experiencing 24 hour oviposition periods.[22] When the number of fruits and hosts were increased by 4 folds, the average number of eggs laid per day increased from about 13 eggs to approximately 24 days.[22] The eggs are laid and will hatch in flavedo part of the fruit.[23]

As a form of egg guarding and preventing overpopulation in a fruit, A. suspensa females have been shown to deposit pheromones from anal membranes that deter repeated attempts of oviposition in a fruit.[20] After extracting and isolating this water-soluble pheromone, it has been learned that it may deter boring attempts by A. suspensa for at least 6 days.[20]

In order to understand the site selection for egg-laying, scientists have observed the ovipositional in wild and laboratory-reared A. suspensa responses when presented an array of chemical stimuli.[22] By noticing that sucrose and organic salts did not appear to influence the oviposition site in either group, it was learned that A. suspensa may not have place much importance in primarily plant constituents in oviposition site.[22] It was discovered that plant compounds such as naringin and quinine were able to inhibit oviposition, reducing the areas in which A. suspensa may inhabit and reproduce.[22] No significant difference was observed between wild and lab-reared A. suspensa for response to oviposition-deterring chemicals.[22]

Social behavior

Social behavior of A. suspensa specifically remains relatively unexplored. The eggs of A. suspensa  may be deposited in groups or as individuals based on the size of the fruit.[23] The larvae inside the eggs will then hatch in the flavedo and all move towards the pulp[disambiguation needed] as they grow. The larvae of A. suspensa may feed on the fruit constantly over a 24 hour period, but no group sociality has been studied apart from feeding on the same areas.[23] Adult sociality has only been studied in males fighting for single leaf territories in order to improve chances of mating.[23] The exploration of cooperation in A. suspensa may be a further area of interest.[vague]

Enemies

For A. suspensa, there are a few natural enemies like P. cindemmaiae and E. annulipes[24]. These predators are classified as parasitoids, or an insect whose larvae live as parasites and end up killing their host. Historically, these parasitoids have been introduced by humans in Puerto Rico and Florida to help combat against A. suspensa[24][8].

Research has emphasized the role of biological control of A. suspensa and less on natural parasites and predators. There have been an array of species like A. indica, D. anastrephilus, and T. daci that infect A. suspensa as larvae. By doing this, they are able to gain from the larvae and pupae of A. suspensa, as the host is harmed.[8]

Due to the large economic damage that A. suspensa may inflict on society, there have been extensive studies on biological control of A. suspensa. An example of this biological control is the endoparasite braconid wasp, D. longicaudata, which indicated a 40% reduction in A. suspensa  populations. The small wasps attack A. suspensa by laying eggs in A. suspensa larvae.[8] When the fly enters the pupal stage, this is when the eggs of the wasp begin to hatch and feed on the larvae. Due to this leeching of resources from A. suspensa, the pupa is often killed before it can develop into an adult.[citation needed]

Earliest studies of control A. suspensa to protect fruits started with ideas as simple as using hot water; now the studies and solutions are much more intricate.[25] Recent studies in Florida have focused on biological control of  A. suspensa with the use of entomopathogenic nematodes.[26] There have also been studies using imidacloprid-treated spheres for control and eradication of A. suspensa in areas where it may be difficult to spread a large range of insecticides.[27]

Mimicry and protective coloration

There is no apparent mimicry seen by A. suspensa or of other insects specifically mimicking A. suspensa. It may be possible that the yellow and brown color may be adopted from bees to discourage predators, but no studies have thoroughly examined this form of mimicry.[2]

Physiology

Studies on A. suspensa physiology have greatly matched the physiology of other species in Anastrepha.[citation needed]

Flight

It has been shown that flight in A. suspensa may be influenced by age, size, sex, and weight. On average, adult A. suspensa can fly 120 kilometers, which allows for the spreading of the species.[2] Studies have examined the effect of gamma irradiation on flies at different stages of their life to make them sterile. It was observed that females that were treated with gamma irradiation fly farther and faster early in their life than untreated females but slower later in their lives.[28] This difference may be due to affecting the female's role in reproduction. For males, being sterilized by gamma irradiation did not produce significant differences.[28] Gamma radiation also changed the waveform of the sound created by the wings of the flies. By sterilizing the pupae, it was observed that wing beat frequency decreased.[28]

Light and Vision

While most studies on A. suspensa have been in a laboratory setting, the results have been telling for the set-up of vision in A. suspensa. Research has shown clear inclinations for A. suspensa towards objects that are colored orange.[29] This preference is then followed by yellow and yellow-green. Colors that did not greatly attract A. suspensa were dark green, red, blue, white, and black.[29] Based on these preferences, it was hypothesized that the photoreceptors in A. suspensa respond specifically to 580-590 nm visible light range. Responses to this light range seems to correspond with fruit seeking. Other studies have discussed the strong sense of vision for A. suspensa and how this has allowed for conducting experiments with clear cages to further test the behavior of the flies.[29]

Sound and Hearing

A. suspensa on a leaf

In the process of creating a better way to search fruits for the presence of A. suspensa larva than cutting the fruit, scientists discovered that A. suspensa larvae and pupae create sounds while chewing.[16] While the interactions based on sound between organisms at this age have not been studies, the sounds are important for scientists in understanding the life cycle in terms of feeding with A. suspensa[16].

Gustation

Since larvae burrow beneath the surface of fruits that they occupy, it is difficult to observe feeding behavior of these early stages of the flies. In adult A. suspensa, it has been noted that food coloring does not affect the fitness parameters of the flies. Additionally, it was learned that A. suspensa ingested both liquids and solid particles. This information elucidated a feeding mechanism similar to that of the Mediterranean fly.[citation needed]

Microbiome

While A. suspensa has not been identified with gut microbiome directly, there are indirect interactions. It has been studied and noted that A. suspensa interact with different bacteria microbial violates. It has been noted specifically in E. agglomerans and other Enterobacteriaceae that A. suspensa females are attracted to their volatile chemicals.[30] Despite this preliminary study, there is not enough evidence to fully support direct ties and interactions between these bacteria and A. suspensa.

Mutualism

Based on research that has been conducted in A. suspensa, there are no clear cases of mutualism with either plants, animals, or microbes. These flies resemble partially parasitic relationship with fruits and they may be influencing the ability of A. suspensa to maintain mutualistic relationships. Even though typically A. suspensa infest spoiled and rotten fruit, they may be influencing the ability for the fruit to disperse its seeds by making it less appealing to animals. At the same time, A. suspensa larva reap the benefits of the fruit for growth.[24][8]

Interactions with humans

Historically, adult A. suspensa are considered the most serious fruit fly pests tropical fruit fly pests. Due to their large array of host fruits ranging from guava to oranges, the economic damage is expansive.[31] A. suspensa are believed to be highly invasive due their potential to their broad native range and strong ability to adapt to different environments.[31] These interactions are amplified by the high likelihood of entry into different countries through legal and illegal means.[31] A. suspensa are also difficult to detect and to control without causing expansive damage, making them very costly combat.[31] In order to inhibit the spread of the flies, many countries like Bermuda, Brazil, China, Colombia, Japan, New Zealand, the Philippines, Thailand, and the United States have adopted strict protocols to maintain fly-free zones when importing fruit.[8]

Conservation

Since A. suspensa is seen as an agricultural and economic threat, there are no cases of conservation of A. suspensa. The parasitic nature of the fly has allowed A. suspensa to expand into various habitats. There is a case on how A. suspensa used to reside in California; however, it no longer is spotted in the Southern California region, indicating unexplained habitat loss.[13][8]

References

  1. ^ Sivinski, J.M.; Calkins, C.O.; Baranowski, R.; Harris, D.; Brambila, J.; Diaz, J.; Burns, R.E.; Holler, T.; Dodson, G. (1996). "Suppression of a Caribbean Fruit Fly (Anastrepha suspensa(Loew) Diptera: Tephritidae) Population through Augmented Releases of the ParasitoidDiachasmimorpha longicaudata(Ashmead) (Hymenoptera: Braconidae)". Biological Control. 6 (2): 177–85. doi:10.1006/bcon.1996.0022.
  2. ^ a b c d e f g h i Stone, Alan (1942). The fruitflies of the genus Anastrepha. U.S. Dept. of Agriculture.
  3. ^ "EPPO Global Database". https. Retrieved 2019-10-01.
  4. ^ a b c Foote, B. A. (1994-05-01). "Handbook of the Fruit Flies (Diptera: Tephritidae)of America North of Mexico". Annals of the Entomological Society of America. 87 (3): 400–401. doi:10.1093/aesa/87.3.400. ISSN 1938-2901.
  5. ^ Dutra, Vivian S.; Ronchi-Teles, Beatriz; Steck, Gary J.; Rodriguez, Erick J.; Norrbom, Allen L.; Sutton, Bruce D.; Silva, Janisete G. (January 2018). "Description of the Larvae of Anastrepha curitis, Anastrepha pickeli and Anastrepha pulchra (Diptera: Tephritidae)". Proceedings of the Entomological Society of Washington. 120 (1): 9–24. doi:10.4289/0013-8797.120.1.9. ISSN 0013-8797.
  6. ^ A. Canal, Nelson; Hernández-Ortiz, Vicente; Tigrero, Juan; Selivon, Denise (2015-11-26). "Morphometric study of third-instar larvae from five morphotypes of the Anastrepha fraterculus cryptic species complex (Diptera, Tephritidae)". ZooKeys (540): 41–59. doi:10.3897/zookeys.540.6012. ISSN 1313-2970. PMID 26798253.{{cite journal}}: CS1 maint: unflagged free DOI (link)
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  12. ^ a b Herández-Ortiz, Vicente; Zucchi, Roberto; Norrbom, Allen (1999-12-20), "Phylogeny of the Genera Anastrepha and Toxotrypana (Trypetinae", Fruit Flies (Tephritidae), CRC Press, pp. 299–342, doi:10.1201/9781420074468.ch12, ISBN 9780849312755
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  14. ^ a b c d e f g h "Caribbean fruit fly - Anastrepha suspensa". entomology.ifas.ufl.edu. Retrieved 2019-10-02.
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  16. ^ a b c d e Calkins, C. O.; Webb, J. C. (1988). "Temporal and Seasonal Differences in Movement of the Caribbean Fruit Fly Larvae in Grapefruit and the Relationship to Detection by Acoustics". The Florida Entomologist. 71 (4): 409–416. doi:10.2307/3495000. ISSN 0015-4040. JSTOR 3495000.
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  19. ^ a b Shelly, Todd E. (2004-09-01). "Scent Marking by Males of the Mediterranean Fruit Fly, Ceratitis capitata (Diptera: Tephritidae)". Journal of Insect Behavior. 17 (5): 709–722. doi:10.1023/B:JOIR.0000042551.10590.d2. ISSN 1572-8889.
  20. ^ a b c Prokopy, Ronald J.; Greany, P. D.; Chambers, D. L. (1977-06-01). "Oviposition-Deterring Pheromone in Anastrepha suspensa". Environmental Entomology. 6 (3): 463–465. doi:10.1093/ee/6.3.463. ISSN 0046-225X.
  21. ^ a b c d e f g h Burk, T.; Webb, J. C. (1983-07-01). "Effect of Male Size on Calling Propensity, Song Parameters, and Mating Success in Caribbean Fruit Flies, Anastrepha suspensa (Loew) (Diptera: Tephritidae)". Annals of the Entomological Society of America. 76 (4): 678–682. doi:10.1093/aesa/76.4.678. ISSN 0013-8746.
  22. ^ a b c d e f Lawrence, Pauline O.; Greany, P. D.; Nation, J. L.; Baranowski, R. M. (1978-03-15). "Oviposition Behavior of Biosteres longicaudatus, a Parasite of the Caribbean Fruit Fly, Anastrepha suspensa". Annals of the Entomological Society of America. 71 (2): 253–256. doi:10.1093/aesa/71.2.253. ISSN 0013-8746.
  23. ^ a b c d Aluja, M (1994). "Bionomics and Management of Anastrepha". Annual Review of Entomology. 39 (1): 155–178. doi:10.1146/annurev.en.39.010194.001103.
  24. ^ a b c Sivinski, J. M.; Calkins, C. O.; Baranowski, R.; Harris, D.; Brambila, J.; Diaz, J.; Burns, R. E.; Holler, T.; Dodson, G. (1996-04-01). "Suppression of a Caribbean Fruit Fly (Anastrepha suspensa(Loew) Diptera: Tephritidae) Population through Augmented Releases of the ParasitoidDiachasmimorpha longicaudata(Ashmead) (Hymenoptera: Braconidae)". Biological Control. 6 (2): 177–185. doi:10.1006/bcon.1996.0022. ISSN 1049-9644.
  25. ^ Sharp, Jennifer L. (1986-06-01). "Hot-water Treatment for Control of Anastrepha suspensa (Diptera: Tephritidae) in Mangos". Journal of Economic Entomology. 79 (3): 706–708. doi:10.1093/jee/79.3.706. ISSN 0022-0493.
  26. ^ Heve, William K.; El-Borai, Fahiem E.; Carrillo, Daniel; Duncan, Larry W. (June 2017). "Biological control potential of entomopathogenic nematodes for management of Caribbean fruit fly, Anastrepha suspensa Loew (Tephritidae)". Pest Management Science. 73 (6): 1220–1228. doi:10.1002/ps.4447. ISSN 1526-4998. PMID 27717178.
  27. ^ Liburd, Oscar E.; Holler, Timothy C.; Moses, Amy L. (April 2004). "Toxicity of imidacloprid-treated spheres to Caribbean fruit fly, Anastrepha suspensa (Diptera: Tephritidae) and its parasitoid Diachasmimorpha longicaudata (Hymenoptera: Braconidae) in the laboratory". Journal of Economic Entomology. 97 (2): 525–529. doi:10.1603/0022-0493-97.2.525. ISSN 0022-0493. PMID 15154477.
  28. ^ a b c Gould, Walter P. (June 1998). "Cold Torpor and Flight Threshold of Anastrepha suspensa (Diptera: Tephritidae)". The Florida Entomologist. 81 (2): 211–216. doi:10.2307/3496088. ISSN 0015-4040. JSTOR 3496088.
  29. ^ a b c Kendra, Paul E.; Montgomery, Wayne S.; Epsky, Nancy D.; Heath, Robert R. (2009-08-01). "Electroantennogram and Behavioral Responses of Anastrepha suspensa (Diptera: Tephritidae) to Putrescine and Ammonium Bicarbonate Lures". Environmental Entomology. 38 (4): 1259–1266. doi:10.1603/022.038.0437. ISSN 0046-225X. PMID 19689908.
  30. ^ Leroy, Pascal D.; Sabri, Ahmed; Verheggen, François J.; Francis, Frédéric; Thonart, Philippe; Haubruge, Eric (2011-09-01). "The semiochemically mediated interactions between bacteria and insects". Chemoecology. 21 (3): 113–122. doi:10.1007/s00049-011-0074-6. ISSN 1423-0445.
  31. ^ a b c d Emden, H. F. (January 1989). "World Crop Pests, Vol. 2. Aphids; their biology, natural enemies and control, PART B (Eds. A. K. Minks and P. Harrewijn). Elsevier, Amsterdam". Entomologia Experimentalis et Applicata. 50 (1): 36. doi:10.1111/j.1570-7458.1989.tb02311.x. ISSN 0013-8703.

Further reading