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''Helicoverpa assulta'', the oriental tobacco budworm, is a
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
of the family
Noctuidae The Noctuidae, commonly known as owlet moths, cutworms or armyworms, are a family of moths. They are considered the most controversial family in the superfamily Noctuoidea because many of the clades are constantly changing, along with the other f ...
. ''H. assulta'' adults are migratory and are found all over the Old World Tropics including Asia, Africa, and Australia. This species has a brown coloured pattern on their
forewing Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindwin ...
s while their
hindwing Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindwi ...
s are yellowish orange and have a brown margin which has a pale mark. The
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan of ...
is about . H. assulta is closely related to
Helicoverpa armigera ''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 lattermost in the UK, where it is a migrant). The larvae f ...
and
Helicoverpa zea ''Helicoverpa zea, ''commonly known as the corn earworm, is a species (formerly in the genus ''Heliothis'') in the family Noctuidae. The larva of the moth ''Helicoverpa zea'' is a major agricultural pest. Since it is polyphagous (feeds on many ...
. The larvae feed on various ''
Solanaceae The Solanaceae , or nightshades, are a family of flowering plants that ranges from annual and perennial herbs to vines, lianas, epiphytes, shrubs, and trees, and includes a number of agricultural crops, medicinal plants, spices, weeds, and orn ...
'' species, including ''
Lycopersicon ''Lycopersicon'' was a genus in the flowering plant family Solanaceae (the nightshades and relatives). It contained about 13 species in the tomato group of nightshades. First removed from the genus ''Solanum'' by Philip Miller in 1754, its remov ...
'', ''
Nicotiana ''Nicotiana'' () is a genus of herbaceous plants and shrubs in the Family (biology), family Solanaceae, that is Native plant, indigenous to the Americas, Australia, Southwestern Africa and the South Pacific. Various ''Nicotiana'' species, common ...
'' (''
Nicotiana tabacum ''Nicotiana tabacum'', or cultivated tobacco, is an annually grown herbaceous plant of the ''Nicotiana'' genus. The plant is tropical in origin, is commonly grown throughout the world, and is often found in cultivation. It grows to heights bet ...
''), ''
Physalis ''Physalis'' (, , , , from φυσαλλίς ''phusallís'' "bladder") is a genus of approximately 75 to 90 flowering plants in the Solanum, nightshade family (biology), family (Solanaceae), which are native to the Americas and Australasia. At l ...
'' and ''
Solanum ''Solanum'' is a large and diverse genus of flowering plants, which include three food crops of high economic importance: the potato, the tomato and the eggplant (aubergine, brinjal). It is the largest genus in the nightshade family Solanaceae ...
''. It also feeds on fruits of ''
Physalis peruviana ''Physalis peruviana'' is a species of plant in the nightshade family (Solanaceae) native to Colombia, Ecuador and Peru. It is commonly known as Cape gooseberry, goldenberry or ground cherry, as poha in Hawaii, and as Harankash in Egypt, while i ...
'' and ''
Datura ''Datura'' is a genus of nine species of highly poisonous, vespertine-flowering plants belonging to the nightshade family Solanaceae. They are commonly known as thornapples or jimsonweeds, but are also known as devil's trumpets (not to be conf ...
'' species. Because of its feeding behavior, ''H. assulta'' is considered a pest of economically viable crops including tomatoes, tobacco, and hot peppers. Due to the pest status, understanding how to control the moth’s behavior is a priority, but the species’ larval eating behavior and growing resistance to insecticide complicate pest control.


Geographic range

''H. assulta'' is distributed across three continents: Asia, Africa and Australia. In Asia, populations of ''H. assulta'' are specifically found in
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
,
Korea Korea ( ko, 한국, or , ) is a peninsular region in East Asia. Since 1945, it has been divided at or near the 38th parallel, with North Korea (Democratic People's Republic of Korea) comprising its northern half and South Korea (Republic o ...
,
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
, and
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
. Environmental conditions, like temperature and geographic locality, can impact the fecundity of the moth.


Food resources

''H. assulta'' is a pest of
red peppers ''Red Peppers'', described as "an interlude with music", is a short comic play in two scenes by Noël Coward. It is one of ten short plays that make up '' Tonight at 8.30'', a cycle written to be performed in groups of three plays across three e ...
(''Capsicum frutescens'' ),
tobacco Tobacco is the common name of several plants in the genus '' Nicotiana'' of the family Solanaceae, and the general term for any product prepared from the cured leaves of these plants. More than 70 species of tobacco are known, but the ...
(Nicotiatna tabacum),
tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...
, and
onion An onion (''Allium cepa'' L., from Latin ''cepa'' meaning "onion"), also known as the bulb onion or common onion, is a vegetable that is the most widely cultivated species of the genus ''Allium''. The shallot is a botanical variety of the onion ...
. They prefer to eat red peppers over tobacco, but are still considered a pest to both. In order to develop and survive successfully, the moth must both have access to nutrient rich food, and be able to digest and uptake the nutrients.


Capsaicin

''H. assulta'' is one of very few insects that can successfully feed on and damage plants, such as hot peppers, containing
capsaicin Capsaicin (8-methyl-''N''-vanillyl-6-nonenamide) ( or ) is an active component of chili peppers, which are plants belonging to the genus ''Capsicum''. It is a chemical irritant for mammals, including humans, and produces a sensation of burning ...
. Studies show that long-term dietary exposure to capsaicin stimulates larger larvae. Furthermore, its unique tolerance to capsaicin may have allowed ''H. assulta'' to expand its host range.


Life Stages


Larvae

Hatching larvae feed on tender or new leaves near the site of oviposition. When it goes through
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
, larvae feed on the fruit and flowers of the host, and can infest the host over the course of their development. Newly hatched larvae exhibit a great preference for tender tobacco leaves, followed by pepper and tomato leaves. The success of ''H. assulta'' is significantly dependent on its larval diet. Larvae that fed on tobacco had a significantly greater fecundity than those that fed on red peppers. The sex ratio differs slightly depending on the type of food the larvae feed on. Larvae who feed on mainly tobacco have a higher female ratio compared to those which feed on mainly pepper. The ''H. assulta'' also preferred tomato as a host plant. The preference for bush red pepper, tobacco, and tomato may be correlated to the semiochemicals or allelochemicals of the host.


Adult

As an adult, the moth prefers to feed on tomatoes during
oviposition 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 typical ...
, followed by tobacco and pepper. Larval diets also impact the number of eggs laid. Adults that previously fed as larvae upon red peppers laid significantly more eggs compared to those that fed on tobacco. Feeding on red peppers also yielded shorter
generation time In population biology and demography, generation time is the average time between two consecutive generations in the lineages of a population. In human populations, generation time typically ranges from 22 to 33 years. Historians sometimes use this ...
, heavier pupae and lower larval and pupal mortality compared to feeding on tobacco.


Mating

Mating occurs during the early hours of the scotophase or the dark phase of a light-dark cycle. However, the mating peak in virgin females tends to advance with age. Unmated adults have also been observed to live longer than mated adults. Prior to mating, females protract and retract the terminal abdomen and vibrate the wings. Mating behavior of adult males includes antennal movement, wing elevation and vibration, extension of hairs, and tapping of the female ovipositor, leading to copulation. Studies also indicate that sex pheromone release is also mediated by PBAN. Female calling and sex pheromone release are also mediated by circadian rhythms entrained to light. This moth is highly sensitive to light intensity, which impacted their sexual behavior. Female calling is inhibited in high-intensity light (50.0 lux), and promoted in low-intensity light (0.5 lux).


Sex Pheromones

Sex pheromones are secreted chemicals used to attract another member of the opposite sex. Because there are different subtypes of H. assulta, each population will respond differently to different ratios and blends of various sex pheromones. Using gas chromatography and mass spectrometry, researchers have identified two sub-populations that respond to different combinations of sex pheromones. There are nine compounds found from female ovipositor washings including hexadecanal, (Z)-9-hexadecenal, (Z)-11-hexadecenal, hexadecyl acetate, (Z)-9-hexadecenyl acetate, (Z)-11-hexadecenyl acetate, hexadecan-l-ol, (Z)-9-hexadecen-l-ol, and (Z)-11-hexadecen-1-ol. The main pheromones that are used are two types of hexadecenals, (Z)-hexadecenal and (Z)-11 hexadecenal, and (Z)-9-hexadecenyl acetate. Field studies conducted in Korea, China, and Thailand also illustrated that a blend of (Z)-9-hexadecenal and (Z)-11-hexadecenal was sufficient for attraction, however the most attractive ratio of compounds varied by specific location. Researchers found that Korean moths preferred a 20:1 blend of (Z)-9-hexadecenal to (Z)-11-hexadecenal. In Thailand, a 7:5:1 blend was most attractive to moths. And In China, the moths found the two different blends were equally attractive.


Release of Pheromones

The suboesophageal ganglion, a portion of the central nervous system in the insect, controls pheromone release. A phermonotropic factor called PBAN (pheromone biosynthesis activating neuropeptide) is synthesized and released into the hemolymph or blood-like fluid found in insects. Because PBAN can be produced independently of the moth's photoperiod, the circadian rhythm of pheromone production must be closely associated with PBAN release. Sex pheromones are only released during scotophase and immediately after pheromone synthesis. Maximum pheromone titer is from day one to day five, and then decreases. The highest concentration of the major female sex pheromone, (Z)-9 hexadecenal (Z9-Z16:Al), is released in a distinct pattern over a 24-hour period or a circadian periodicity. This pattern is affected by age,
photoperiod Photoperiodism is the physiological reaction of organisms to the length of night or a dark period. It occurs in plants and animals. Plant photoperiodism can also be defined as the developmental responses of plants to the relative lengths of light a ...
, and temperature. The daily rhythm of hormone production varies when the female moths are reared under 12 hours of light and 12 hours of darkness vs. continuous light conditions. The maximum pheromone titers in the gland corresponded to the peak calling activities, indicating that these events are synchronous and predictable. In constant darkness, these events were also synchronous. However, when placed in constant light, the H. assulta has two different patterns for pheromone release and calling behavior. Calling was suppressed, but the pheromone release was not. This suggests that these two behaviors are controlled by two different systems, and can somehow communicate to synchronize under normal conditions. When moths are placed in constant light, there is a longer retention and slower decrease of Z9-16: AL in the pheromone gland. The moths may continue to produce the pheromone for a longer period of time, or that the degradation mechanism is inactive and the chemical may only decrease through release. The production of Z9-16: AL is highest under darkness and low-intensity light (0.5 lux), and inhibited at higher-intensity light (5.0 and 50.0 lux).


Pest control

''H. assulta'' is considered to be one of the most destructive pests for various economic crops like tobacco and hot peppers. The moth has caused considerable economic damage to crops in Korea and China. The larvae usually feed on the fruit of host plants. Unlike other insects, it has a high tolerance for the toxicity of capsaicin. While conventional chemical insecticides are necessary for the control of these moths, the growing resistance of the moth and the detrimental effects of overusing these chemicals have drawn researchers to develop other methods of control. Furthermore, the larvae feed inside the fruit, and are therefore protected from the chemical sprays. Studies have illustrated that the moth’s response to insecticide depends on the type of plant it's inhabiting. ''H. assulta'' that were fed on red peppers were more susceptible to certain insecticides such as
fenvalerate Fenvalerate is a synthetic pyrethroid insecticide. It is a mixture of four optical isomers which have different insecticidal activities. The 2-S ''alpha'' (or SS) configuration, known as esfenvalerate, is the most insecticidally active isomer. F ...
, but became more resistant to chemicals like inoxacarb,
phoxim Phoxim is an organophosphate insecticide that is produced by the Bayer corporation. It is an analogous dimethyl ester and an organothiophosphate acaricide. It is allowed for use in limited applications in the European Union. It is banned for ...
and
methomyl Methomyl is a carbamate insecticide introduced in 1966. It is highly toxic to humans, livestock, pets, and wildlife. The EU and UK imposed a pesticide residue limit of 20 µg/kg for apples and oranges. Methomyl is a common active ingredient in c ...
. Current research explores sex pheromone-mediated communication, a method of control aiming to interrupt moth communication. Also, there is a need to understand the complex interactions among the insect, plant, and insecticide.


References


External links


Moths of BorneoAustralian Faunal Driectory
{{Taxonbar, from=Q3342612 assulta Moths described in 1852 Moths of Africa Moths of Asia Moths of Japan Moths of the Middle East Insects of Cape Verde Insects of Chad Insects of the Republic of the Congo Insects of West Africa