Tuta absoluta | |
---|---|
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Lepidoptera |
Family: | Gelechiidae |
Genus: | Tuta |
Species: | T. absoluta |
Binomial name | |
Tuta absoluta | |
Synonyms | |
Tuta absoluta or Phthorimaea absoluta is a species of moth in family Gelechiidae known by the common names South American tomato pinworm, tomato leafminer, tomato pinworm and South American tomato moth. It is well known as a serious pest of tomato crops in Europe, Africa, western Asia and South and Central America, with larvae causing up to 100% loss if not effectively controlled. [2] : 241
T. absoluta was originally described in 1917 by Edward Meyrick as Phthorimaea absoluta, based on individuals collected from Huancayo (Peru). [2] : 240 Later, the pest was reported as Gnorimoschema absoluta, [3] Scrobipalpula absoluta (Povolný), [2] : 240 or Scrobipalpuloides absoluta (Povolný), [2] : 240 but was finally described under the genus Tuta as T. absoluta by Povolný in 1994.< [4] [5] [2] : 240 [6] : 1330
The larva feeds voraciously upon tomato plants, producing large galleries in leaves, burrowing in stalks, and consuming apical buds and green and ripe fruits. It is capable of causing a yield loss of 100%. [7] [2] : 241 Prefers 30 °C (86 °F), requires 14 to 34.6 °C (57.2 to 94.3 °F) for full lifecycle. [2] : 241 Nonetheless cold tolerance does allow for 50% survival of larvae, pupae, and adults, at 0 °C (32 °F). [2] : 241
Its life-cycle comprises four development stages: egg, larva, pupa and adult; combined, 26–75 days. [2] : 241 Adults usually lay yellow [2] : 241 eggs on the underside of leaves or stems, and to a lesser extent on fruits. After hatching, young larvae penetrate leaves, aerial fruits (like tomato) or stems, on which they feed and develop. Pupae (length: 5–6 millimetres (13⁄64–15⁄64 in)) are cylindrical in shape and greenish when just formed becoming darker in color as they are near adult emergence. The pest mainly presents nocturnal habits, and adults usually remain hidden during the day, showing greater morning-crepuscular activity with adults dispersing among crops by flying. Among a range of species within the Solanaceae, tomatoes (Lycopersicon esculentum Miller) appear to be the primary host of T. absoluta.
No evidence of short-day diapause. [2] : 241 Up to 10 generations per year. [2] : 241 Sex pheromone variation was shown by Dominguez et al 2019 to be influenced by the aging process and by plant volatiles. [8]
Adults are 6–7 millimetres (15⁄64–9⁄32 in) in length and present filiform antennae and silver to grey scales. [9] Black spots are present on anterior wings, and the females are wider and more voluminous than the males.
The adult moth has a wingspan around 1 centimetre (3⁄8 in). In favorable weather conditions eight to ten generations can occur in a single year.
Tomato is the main host plant, but T. absoluta also attacks other crop plants of the nightshade family, including potato, [2] : 240 eggplant, pepino, pepper and tobacco. [10] It is known from many solanaceous weeds, including Datura stramonium [11] and Solanum nigrum . [2] : 240
Also known from non-Solanaceae hosts in the Amaranthaceae, Convolvulaceae, Fabaceae, and Malvaceae. [2] : 240
Laboratory rearing is abnormally difficult because T. absoluta requires maternal leaf contact with a suitable host plant for oviposition. [2] : 240–241
This moth was first known as a tomato pest in many South American countries (and Easter Island) [6] : 1330 and was recognized to threaten cultivation in Europe. However, the EU did not list it as an inspection or quarantine pest, and this likely contributed to what happened next. [2] : 241 In 2006, it was identified in Spain [12] [2] : 242, Fig1a from a Chilean parental population introduced to Spain in the early 2000s. [13] [2] : 241 The following year it was detected in France, Italy, Greece, Malta, Algeria and Libya. Morocco in 2008. [2] : 242, Fig1a Starting in 2009, seeing the results of inaction in Europe, the North American Plant Protection Organization, the United States, California, Florida, Canada, and Australia began inspections and preparation for quarantines. [2] : 243 Also in 2009 it was first reported from Turkey. The advance of T. absoluta continued to the east to reach Syria, Lebanon, Jordan, Israel, Iraq and Iran. Further advances southward reached Saudi Arabia, Yemen, Oman and the rest of the Persian Gulf states. In Africa, T. absoluta moved from Egypt to reach Sudan, South Sudan and Ethiopia from the east and to reach the Senegal from the west. It was reported in Nigeria and Zambia [14] and South Africa in 2016. [2] : 242, Fig1a [6] : 1331 An up-to-date global distribution map is available on the Tuta absoluta information network. Reached India and the Himalayans, unconfirmed but possibly also Pakistan and Tajikistan, by 2017. [6] : 1331 In India, Maharashtra state tomato cultivation more affect in November 2016.[ citation needed ] It is now severely infested in Myanmar, especially in tropical tomato growing areas such as Mandalay, Sagaing, Monywa.( April, 2017) In the last few years Tuta absoluta has spread to Kenya. [15] [16] Although it is not there yet, researchers at the University of Guam are concerned about the possible spread of T. absoluta to Guam. [17] As of 2017 [update] USDA's Animal and Plant Health Inspection Service assumed T. absoluta to be present in most of sub-Saharan Africa. [2] : 242 Present in Cape Verde [2] : 243 and Turkey [18] since the 2010-11 survey. [2] : 242, Fig1a
There is a high risk of further invasion northward into more of Central America, and into the United States (a certainty, if it reaches as far as Mexico); [2] : 243 all suitable areas of sub-Saharan Africa [2] : 242 [2] : 242 [2] : 250 and southern Asia; and Australia and New Zealand. [2] : 243 There is a lower risk of invasion in colder areas like Canada, northern Europe, and most of the Russian Federation. [2] : 243
This rapid spread across Mediterranean Europe was due to insufficient coordinated plant protection activity against invasive agricultural pests. [2] : 241
In 2014 the People's Republic of China's Chinese Academy of Agricultural Sciences' Department of Biological Invasions began surveillance and treatment [19] of their own [2] : Sup7, Fig1 [2] : Sup8, Fig2 [20] and neighboring countries (including India and Pakistan) that already have the pest. Surveillance occurs in production areas and near international airports. [2] : 243–4
Losses on tomatoes can reach 100% due to larval feeding, if not effectively controlled. [2] : 241 Even if not that severe, damage will require postharvest inspection expenditures and some financial loss due to unattractive fruit. [2] : 241 The initial European invasion increased tomato production costs by more than 450€/hectare. [2] : 243
Some populations of T. absoluta have developed resistance to organophosphate and pyrethroid pesticides. [21] Newer compounds such as spinosad, [22] imidacloprid [ citation needed ], and Bacillus thuringiensis [23] have demonstrated some efficacy in controlling European outbreaks of this moth. Insecticide costs have increased rapidly, and even that has not always produced good results, due to high quantity application of insecticides that are not especially effective against T. absoluta. As a result, new registrations have been obtained specifically for this pest starting in 2009. Between 2009 and 2011 there was a dramatic increase in authorized APIs and MoAs in Spain and Tunisia for this reason. [6] : 1332–3
A large number of insecticide MoAs are effective, and various ones are registered in various jurisdictions, including: Acetylcholinesterase inhibitors (IRAC group 1B), voltage-gated sodium channel modulators (3A), nicotinic acetylcholine receptor modulators (5), chloride channel activators (6), midgut membrane disruptor (11), oxidative phosphorylation uncouplers (13), nicotinic acetylcholine receptor blockers (15), ecdysone receptor agonists (18), volgate-gated sodium channel blockers (22A and 22B), ryanodine receptor modulators (28), and azadirachtin (of unknown action, UN).
Experiments have revealed some promising agents of biological pest control for this moth, including Nabis pseudoferus , a species of damsel bug, [24] Bacillus thuringiensis , [24] [6] : 1330 : 1332 and Beauveria bassiana . [6] : 1332 Companion planting with Fagopyrum esculentum works by increasing numbers of the parasitoid Necremnus tutae . [2] : 8, Fig2
Relatively natural chemical controls include limonene and borax. [6] : 1332
The sex pheromone for T. absoluta has been identified by researchers at Cornell University and has been found to be highly attractive to male moths. [26] Pheromone lures are used extensively throughout Europe, South America, North Africa and the Middle East for the monitoring and mass-trapping of T. absoluta. The use of pheromone products in combination with a yellow delta trap has been recorded in South Africa. This concept is used to monitor populations of T. absoluta in tomato orchards. [25]
The combined use of pheromones as well as specific light frequency proved to be effective in suppressing the T. absoluta population and keeping it within the economic threshold as it disclosed by Russell IPM in a United Kingdom patent. [27]
Also the use of electric mosquito traps give good results. [28]
Organophosphate and pyrethroid resistance developed in Chile, then in Brazil and (as noted above) Argentina. [21] Spinosad resistance was also first noticed in Chile (possibly thanks to a cytochrome P450 and esterases), and then spinosad/spinetoram cross-resistance in Brazil due to two desensitizing mutations at the same target site: G275E, and an exon-skipping mutation; and perhaps synergistically with other factors. [6] : 1332, T1
Then came the Spanish detection in 2006. The biotype of this European invasion already carried at least 4 resistance mutations from a Chilean [13] [6] : 1331 : 1332–3 : 1334 parental population: 3 in the relevant sodium channel for pyrethroids, [13] including L1014F; [6] : 1332, T1 : 1333 and 1 (A201S) in the enzyme targeted by organophosphates. [29] [6] : 1333
Previously there had been little interest in this subject. Then about six years after the beginning of its invasion of Europe, there was a sharp increase in scientific recognition of - and interest in - resistance in T. absoluta, which only continued to build further year after year. [6] : 1333 : 1334, Fig1 : 1338 In Aydın, Turkey in 2015, the T. absoluta population was found to be highly resistant to indoxacarb, spinosad, chlorantraniliprole, and metaflumizone, but not azadirachtin - while the Urla, İzmir population was only resistant to azadirachtin, and even then only weakly so. [18] Many modes of action have fallen in efficacy in South America and Europe, closely in tandem with popularity of use of those MoAs/insecticides: Abamectin, cartap, indoxacarb, chitin biosynthesis inhibitors, spinosad, and the diamides. Only pyrethroid resistance has been confirmed to have declined. Only chlorfenapyr and Bt toxin have remained at low resistance, likely due to low usage. IRAC (the Insecticide Resistance Action Committee)'s efforts to slow resistance development and spread have been effective in Brazil and Spain, by way of widely disseminated information campaigns targeting the agricultural industries in the area. [6] : Abs : 1336–7 : 1336, Fig2
It has been hypothesized that the flatness of the Brazilian savannah may be speeding up the spread of resistance alleles. [6] : 1331
Interactions between T. absoluta, Bemisia tabaci , resistance, and Neoleucinodes elegantalis , and natural enemies of these pests remain underexplored. There are substantial gaps in knowledge that will need to be filled in the future. [6] : 1338
The first report of diamide/ryanoid (IRAC group 28) failure was in 2015, [30] [6] : 1331 and two years later a related team found this was occurring due to an altered target site due to the mutations G4903E and I4746 M. (These two mutations are parallels of two mutations found to be producing the same results in Plutella xylostella .) Altered binding affinity was found for the mutations G4903 V and I4746T, and they were found in a few resistant populations. In extreme (heterozygotic for resistant alleles) cases the normal application rate becomes hormetic. [31] (The use of chlorantraniliprole for T. absoluta has also resulted in resistance in B. tabaci, even though it is not used against that species, merely because they co-occur on tomato. This is expected to make cyantraniliprole unusable if needed on B. tabaci, in the same area.) [32] [6] : 1338
Resistance to indoxacarb (IRAC group 22A) has appeared due to the mutations F1845Y and V1848I, but is not yet reported for another voltage-dependent sodium channel blocker, metafumizone (22B). (These two mutations, as with the diamides above, have P. xylostella analogues, but in this case these analogues are known to be effective against both indoxacarb and metafumizone.) [6] : 1332, T1 : 1335
Cartap, a nicotinic acetylcholine receptor channel blocker (IRAC group 14), began to show low to moderate efficacy decline in South America starting in 2000, and increasing through at least 2016. Some of this is due to elevated cytochrome P450 activity (see below) possibly as part of demethylation and sulfoxidation detoxification, while less is thought to be due to esterases and glutathione S-transferases. [6] : 1334 (The use of cartap for T. absoluta has also resulted in resistance in B. tabaci, even though it is not used on that species, merely because they co-occur on tomato.) [32] [6] : 1338
Cytochrome P450s are used to resist:
but overall, specific information is still lacking connecting which particular P450s and which particular resistances. [6] : 1334
Bacillus thuringiensis is a gram-positive, soil-dwelling bacterium, the most commonly used biological pesticide worldwide. B. thuringiensis also occurs naturally in the gut of caterpillars of various types of moths and butterflies, as well on leaf surfaces, aquatic environments, animal feces, insect-rich environments, flour mills and grain-storage facilities. It has also been observed to parasitize moths such as Cadra calidella—in laboratory experiments working with C. calidella, many of the moths were diseased due to this parasite.
Insecticides are pesticides used to kill insects. They include ovicides and larvicides used against insect eggs and larvae, respectively. Acaricides, which kill mites and ticks, are not strictly insecticides, but are usually classified together with insecticides. The major use of Insecticides is agriculture, but they are also used in home and garden, industrial buildings, vector control and control of insect parasites of animals and humans. Insecticides are claimed to be a major factor behind the increase in the 20th-century's agricultural productivity. Nearly all insecticides have the potential to significantly alter ecosystems; many are toxic to humans and/or animals; some become concentrated as they spread along the food chain.
Pesticide resistance describes the decreased susceptibility of a pest population to a pesticide that was previously effective at controlling the pest. Pest species evolve pesticide resistance via natural selection: the most resistant specimens survive and pass on their acquired heritable changes traits to their offspring. If a pest has resistance then that will reduce the pesticide's efficacy – efficacy and resistance are inversely related.
Whiteflies are Hemipterans that typically feed on the undersides of plant leaves. They comprise the family Aleyrodidae, the only family in the superfamily Aleyrodoidea. More than 1550 species have been described.
A pyrethroid is an organic compound similar to the natural pyrethrins, which are produced by the flowers of pyrethrums. Pyrethroids are used as commercial and household insecticides.
The Gelechiidae are a family of moths commonly referred to as twirler moths or gelechiid moths. They are the namesake family of the huge and little-studied superfamily Gelechioidea, and the family's taxonomy has been subject to considerable dispute. These are generally very small moths with narrow, fringed wings. The larvae of most species feed internally on various parts of their host plants, sometimes causing galls. Douglas-fir (Pseudotsuga) is a host plant common to many species of the family, particularly of the genus Chionodes, which as a result is more diverse in North America than usual for Gelechioidea.
The silverleaf whitefly is one of several species of whitefly that are currently important agricultural pests. A review in 2011 concluded that the silverleaf whitefly is actually a species complex containing at least 40 morphologically indistinguishable species.
The cabbage looper is a medium-sized moth in the family Noctuidae, a family commonly referred to as owlet moths. Its common name comes from its preferred host plants and distinctive crawling behavior. Cruciferous vegetables, such as cabbage, bok choy, and broccoli, are its main host plant; hence, the reference to cabbage in its common name. The larva is called a looper because it arches its back into a loop when it crawls.
Deltamethrin is a pyrethroid ester insecticide. Deltamethrin plays a key role in controlling malaria vectors, and is used in the manufacture of long-lasting insecticidal mosquito nets; however, resistance of mosquitos and bed bugs to deltamethrin has seen a widespread increase.
A leaf miner is any one of numerous species of insects in which the larval stage lives in, and eats, the leaf tissue of plants. The vast majority of leaf-mining insects are moths (Lepidoptera), sawflies, and flies (Diptera). Some beetles also exhibit this behavior.
Helicoverpa armigera is a species of Lepidoptera in the family Noctuidae. It is known as the cotton bollworm, corn earworm, Old World (African) bollworm, or scarce bordered straw. The larvae feed on a wide range of plants, including many important cultivated crops. It is a major pest in cotton and one of the most polyphagous and cosmopolitan pest species. It should not be confused with the similarly named larva of the related species Helicoverpa zea.
Spinosad is an insecticide based on chemical compounds found in the bacterial species Saccharopolyspora spinosa. The genus Saccharopolyspora was discovered in 1985 in isolates from crushed sugarcane. The bacteria produce yellowish-pink aerial hyphae, with bead-like chains of spores enclosed in a characteristic hairy sheath. This genus is defined as aerobic, Gram-positive, nonacid-fast actinomycetes with fragmenting substrate mycelium. S. spinosa was isolated from soil collected inside a nonoperational sugar mill rum still in the Virgin Islands. Spinosad is a mixture of chemical compounds in the spinosyn family that has a generalized structure consisting of a unique tetracyclic ring system attached to an amino sugar (D-forosamine) and a neutral sugar (tri-Ο-methyl-L-rhamnose). Spinosad is relatively nonpolar and not easily dissolved in water.
Cyhalothrin is the ISO common name for an organic compound that, in specific isomeric forms, is used as a pesticide. It is a pyrethroid, a class of synthetic insecticides that mimic the structure and properties of the naturally occurring insecticide pyrethrin which is present in the flowers of Chrysanthemum cinerariifolium. Pyrethroids such as cyhalothrin are often preferred as an active ingredient in agricultural insecticides because they are more cost-effective and longer acting than natural pyrethrins. λ-and γ-cyhalothrin are now used to control insects and spider mites in crops including cotton, cereals, potatoes and vegetables.
Chloridea virescens, commonly known as the tobacco budworm, is a moth of the family Noctuidae found throughout the eastern and southwestern United States along with parts of Central America and South America.
Helicoverpa assulta, the oriental tobacco budworm, is a moth of the family Noctuidae. H. assulta adults are migratory and are found all over the Old World Tropics including Asia, Africa, and Australia.
The South American potato tuber moth, Andean potato tuber moth or tomato stemborer is a moth of the family Gelechiidae. It is native to South America, but has become a pest worldwide. Records include North America, Australia and New Zealand.
Tefluthrin is the ISO common name for an organic compound that is used as a pesticide. It is a pyrethroid, a class of synthetic insecticides that mimic the structure and properties of the naturally occurring insecticide pyrethrin which is present in the flowers of Chrysanthemum cinerariifolium. Pyrethroids such as tefluthrin are often preferred as active ingredients in agricultural insecticides because they are more cost-effective and longer acting than natural pyrethrins. It is effective against soil pests because it can move as a vapour without irreversibly binding to soil particles: in this respect it differs from most other pyrethroids.
Cyantraniliprole is an insecticide of the ryanoid class, specifically a diamide insecticide. It is approved for use in the United States, Canada, China, and India. Because of its uncommon mechanism of action as a ryanoid, it has activity against pests such as Diaphorina citri that have developed resistance to other classes of insecticides.
Macrolophus pygmaeus is a species of plant bug in the family Miridae. It is found in Europe except the high north, south to north Africa and east to Asia Minor then to Central Asia. This species is omnivorous, preying on Tuta absoluta eggs and larvae, Ephestia kuehniella eggs, Macrosiphum euphorbiae nymphs, and plants such as Vicia fava. When feeding on plants, M. pygmaeus consumes extrafloral nectar. Its varied diet has created interest in M. pygmaeus as a pest control insect for the prior mentioned species.
Chilli leaf curl virus(ChiLCV) is a DNA virus from the genus Begomovirus and the family Geminiviridae. ChiLCV causes severe disease especially in pepper (Capsicum spp.), but also affects other crops such as tomato (Solanum lycopersicum). It can be found in tropical and subtropical regions primarily in India, but has also been detected in countries such as Indonesia and Sri Lanka. This virus is transmitted by an insect vector from the family Aleyrodidae and order Hemiptera, the whitefly Bemisia tabaci. The primary host for ChiLCV are several Capsicum spp., but host species also include tomato and amaranth. ChiLCV has been responsible for several epidemics and causes severe economic losses. It is the focus of research trying to understand the genetic basis of resistance. Currently, a few sources of resistance have been discovered and used to breed resistant varieties.
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