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''Agrotis ipsilon'', the dark sword-grass, black cutworm, greasy cutworm, floodplain cutworm or ipsilon dart, is a small
noctuid 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 ...
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 ...
found worldwide. The moth gets its scientific name from black markings on its forewings shaped like the letter "Y" or the Greek letter
upsilon Upsilon (, ; uppercase Υ, lowercase υ; el, ''ýpsilon'' ) or ypsilon is the 20th letter of the Greek alphabet. In the system of Greek numerals, grc, Υʹ, label=none has a value of 400. It is derived from the Phoenician waw . Etym ...
. The larvae are known as "
cutworms Cutworms are moth larvae that hide under litter or soil during the day, coming out in the dark to feed on plants. A larva typically attacks the first part of the plant it encounters, namely the stem, often of a seedling, and consequently cuts it ...
" because they cut plants and other crops. The larvae are serious agricultural
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
and feed on nearly all varieties of
vegetable Vegetables are parts of plants that are consumed by humans or other animals as food. The original meaning is still commonly used and is applied to plants collectively to refer to all edible plant matter, including the flowers, fruits, stems, ...
s and many important
grain A grain is a small, hard, dry fruit (caryopsis) – with or without an attached hull layer – harvested for human or animal consumption. A grain crop is a grain-producing plant. The two main types of commercial grain crops are cereals and legum ...
s.Capinera, John L
"Common Name: Black Cutworm"
''Entomology and Nematology'', University of Florida, October 2006.
This species is a seasonal migrant that travels north in the spring and south in the fall to escape extreme temperatures in the summer and winter. The migration patterns reflect how reproduction occurs in the spring and ceases in the fall. Females release sex pheromones to attract males for mating. Pheromone production and release in females and pheromone responsiveness in males is dependent on the juvenile hormone (JH) and pheromone biosynthesis activating neuropeptide (BPAN). In the span of 2 months, the moth progresses through the life cycle stages egg, larvae, pupa, and adult. Throughout this time period, this moth faces the risk of predation and parasitism, such as by ''Hexamermis arvalis'' or by the parasite ''Archytas cirphis''.


Description

38–48 mm. Antennae in male bipectinated. Forewings brown, reddish-tinged, mixed with pale greyish-ochreous, costa and sometimes median area suffused with dark fuscous ; first and second lines edged with dark fuscous; spots outlined with black, reniform followed by a short black dash ; subterminal line obscure, usually preceded in middle by two fine black marks. Hindwings whitish-grey or whitish, terminally suffused with fuscous. Larva ochreous brownish or bronzy-grey, sides sometimes greenish ; dorsal, subdorsal, and spiracular lines faintly darker or lighter, usually darker-edged ; head sufFusedly brown-marked.


Geographic range

Populations of this species have been found in southern
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
, 48 of the United States (and additionally
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
),
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
, Central and South America,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
,
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
, the
Pacific Rim The Pacific Rim comprises the lands around the rim of the Pacific Ocean. The ''Pacific Basin'' includes the Pacific Rim and the islands in the Pacific Ocean. The Pacific Rim roughly overlaps with the geologic Pacific Ring of Fire. List of co ...
,
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
, Europe, and Asia.Showers, William B. "Migratory Ecology of the Black Cutworm." ''Annual Review of Entomology'', vol. 42, 1997, pp. 393–425. However, they are absent from some tropical regions and colder areas and are more widespread in the Northern than Southern Hemisphere. This species is also known to migrate north in the spring and migrate south in the fall.


Food resources


Caterpillars

Larvae feed on weeds such as bluegrass,
curled dock ''Rumex crispus'', the curly dock, curled dock or yellow dock, is a perennial flowering plant in the family Polygonaceae, native to Europe and Western Asia. Description The plant produces an inflorescence or flower stalk that grows to high. I ...
,
lambsquarters Lamb's quarter, lambsquarters, and similar terms refer to any of various edible species of herbaceous plants otherwise known by the common names goosefoot or pigweed. There are numerous variations, with or without hyphens and apostrophes, using on ...
,
yellow rocket ''Barbarea'' (winter cress or yellow rocket) is a genus of about 22 species of flowering plants in the family Brassicaceae, native to temperate regions of the Northern Hemisphere, with the highest species diversity in southern Europe and southwe ...
, and
redroot pigweed ''Amaranthus retroflexus'' is a species of flowering plant in the family Amaranthaceae with several common names, including red-root amaranth, redroot pigweed, red-rooted pigweed, common amaranth, pigweed amaranth, and common tumbleweed. page 47 ...
. They will often eat all the weeds available before moving to attacking crops. Favored crops include most vegetable plants, alfalfa, clover, cotton, rice, sorghum, strawberry, sugarbeet, tobacco, and occasionally grains and grasses.


Adults

Adults feed on flower
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
. They are also attracted to deciduous trees and shrubs such as linden, wild plum,
crabapple ''Malus'' ( or ) is a genus of about 30–55 species of small deciduous trees or shrubs in the family Rosaceae, including the domesticated orchard apple, crab apples, wild apples, and rainberries. The genus is native plant, native to the temper ...
, and
lilac ''Syringa'' is a genus of 12 currently recognized species of flowering plant, flowering woody plants in the olive family or Oleaceae called lilacs. These lilacs are native to woodland and scrub from southeastern Europe to eastern Asia, and wid ...
. They are a pollinator of fetterbush lyonia.


Parental care


Oviposition

Based on the types of debris on the ground, the black cutworm prefers to oviposit in areas with fencerow (pasture) debris rather than corn field debris, woodland floor debris, and bare soil. Fencerow debris includes dry grass debris, and this may be attractive for females to oviposit early in the spring before rapid vegetation growth occurs. After this growth, though, the moths are drawn more to low, dense plants such as the
curled dock ''Rumex crispus'', the curly dock, curled dock or yellow dock, is a perennial flowering plant in the family Polygonaceae, native to Europe and Western Asia. Description The plant produces an inflorescence or flower stalk that grows to high. I ...
and
yellow rocket ''Barbarea'' (winter cress or yellow rocket) is a genus of about 22 species of flowering plants in the family Brassicaceae, native to temperate regions of the Northern Hemisphere, with the highest species diversity in southern Europe and southwe ...
. These plants have multiple stems and many low-lying basal leaves. On most plant species, the cutworm prefers to oviposit on the leaves rather than the stem.


Life history


Life cycle

In a given year, the number of generations differs based on location and weather conditions. In Canada, there are 1 or 2 generations, while in the United States, there are 2 to 4 per year. This species is abundant in warmer temperatures (such as Arkansas, US) during the late spring in May–June and early fall in September and October, while they are more abundant in cooler temperature (such as New York, US) during the summer in June and July. One life cycle lasts between 35–60 days.


Egg

The egg stage lasts 3 to 6 days. Females oviposit eggs in clusters on low-lying leaves. If such host plants are not available, the females will oviposit on dead plant material. However, they will not lay eggs on bare soil. Females can deposit eggs singly, or in groups of up to 1200 to 1900 eggs. The nearly spherical eggs are initially white but turn brown with age. The surface of the egg possesses 35–40 ribs that radiate from one apex.


Caterpillar

The larval stage lasts 20–40 days. Over the span of 5 to 9
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 ...
s, the caterpillar body grows from 3.5 mm to a maximum of 55 mm. Larval development is optimized at a temperature of 27 degrees Celsius, and instars 1–5 are most successful at higher humidities. By the 4th instar, the larva becomes light sensitive and spends most of the daylight underground. The larva are considered pests because they damage the plant tissue under the soil. The larvae are cannibalistic. The larva can range in color from light gray or gray brown to black. The ventral side is usually lighter, and this species does not have a dorsal band. The entire body is covered with granules and the head possesses many dark spots.


Pupa

The pupal stage lasts 12–20 days. This species pupates under the soil approximately 3–12 mm below the surface. The pupae appear to be dark brown and are 17–12 mm long and 5–6 mm wide.


Adult

One complete generation from egg to adult lasts 35–60 days. The female preoviposition period lasts 7–10 days. Adults have a wingspan of 40–55 mm. The forewings are dark brown, and the distal area has a light irregular band a black dash mark. The hindwings are whitish to gray and have darker colored veins.


Nature Of Damage

Gram cutworms are active in the Rabi season and damage the crop. * In India, winter is generally the lean season for insects and pests. But cutworms belong to that small insect group. Those who damage the
Rabi crop Rabi crops or rabi harvest, also known as winter crops, are agricultural crops that are sown in winter and harvested in the spring in India, Pakistan and Bangladesh. The complimentary of the rabi crop is the kharif crop, which is grown after t ...
by their destructive activity. * cutworms cut the plant down into the ground or twist the whole plant like a gram, hence it is known as a cutworm. * cutworm is also known as a surface caterpillar because it is a sai surface and performs its activities a few centimetres. * The larva of Gram cutworm causes great damage to the crop. Its
larvae A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
disappear during the day and come out in the evening to damage the crop. * First of all, they feed on the epidermis of the fallen leaves or green leaves touching the ground letter
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
cut the leaf, shoot the plant just above the ground level, and buried in the soil. * cutworm damages many plants causing serious damage. It attacks the gram field between November and February.


Migration

''A. ipsilon'' are seasonal migratory insects that travel south in the fall to escape harsh cold temperatures and travel north in the spring to escape extremely warm weather. Therefore, changes in thermoperiod as well as
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 ...
may influence the onset of migration patterns in this species. Before migration southward in the fall, the reproductive system in both females and males shuts down to prevent
copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetrat ...
before winter. In the spring and early summer, though, before migration north, females release sex pheromones soon after eclosion. In one study, female moths collected from late April to early May were 100% mated.


Enemies


Predators

Several species of
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s prey on the black cutworm. Larvae parasitized by ''
Meteorus leviventris ''Meteorus rubens'' is a species of parasitoid wasp in the family Braconidae The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with a ...
,'' a type of parasitoid, eat 24% less vegetation and cut 36% fewer seedlings. Other
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s include several fly species such as '' Archytas cirphis'', ''
Bonnetia comta ''Bonnetia'' is a genus of flowering plants in the family Bonnetiaceae. Most of the roughly 30 species are shrubs. The remaining species, all trees, are among the dominant species in the forest vegetation on the tepui plateaus of northern South A ...
'', ''
Eucelatoria armigera ''Eucelatoria'' is a genus of flies in the family Tachinidae. Species *'' Eucelatoria albopilosa'' ( Curran, 1926) *'' Eucelatoria argentea'' (Thompson, 1968) *'' Eucelatoria armigera'' ( Coquillett, 1889) *'' Eucelatoria aurata'' (Townsend, 192 ...
'' and ''
Sisyropa eudryae ''Sisyropa'' is a genus of flies in the family Tachinidae. Species *'' S. alypiae'' Sellers, 1943 *'' S. eudryae'' (Townsend, 1892) *'' S. formosa'' Mesnil, 1944 *'' S. heterusiae'' (Coquillett, 1899) *'' S. picta'' (Baranov, 1935) *'' S. promin ...
.''
Ground beetle Ground beetles are a large, cosmopolitan distribution, cosmopolitan family (biology), family of beetles, the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 in Europe. As of 2015, it i ...
s also eat black cutworm larvae. Ants, specifically ''
Lasius neoniger ''Lasius neoniger'', also known as the turfgrass ant, Labour day ant, cornfield ant or nuisance ant, is a species of ant in the genus ''Lasius''. Found in North America, the species is common in the eastern United States and Canada, though they ...
'' also prey on this species and feed on ''A. ipsilon'' eggs''.''López, Rolando, and Daniel A. Potter. "Ant Predation on Eggs and Larvae of the Black Cutworm (Lepidoptera: Noctuidae) and Japanese Beetle (Coleoptera: Scarabaeidae) in Turfgrass." ''Environmental Entomology'', vol. 29, no. 1, Jan. 2000, pp. 116–125., doi:10.1603/0046-225x-29.1.116.


Parasites

An entomopathogenic nematode called ''Hexamermis arvalis'' is known to infect 60% of larvae in the central United States. This parasite ultimately kills the insect. The parasite thrives in moist soil conditions.


Mating


Female calling behavior

Calling behavior is the act of females releasing sex pheromones in preparation for mating. Calling behavior increases within the first three days after eclosion but decreases as the females grow older. As well, as the females grow older, they onset time of calling behavior occurs earlier. Calling earlier allows older females to have increased mating success as they normally produce less sex pheromone and need to appear more attractive than younger females. The amount of sex pheromone in the body and calling behavior are coordinated on a time scale.


Pheromone biosynthesis activating neuropeptide

Females produce a sex pheromone in the pheromone gland on their abdominal tips that attracts males for mating. Biosynthesis of the sex pheromone is controlled by a
neurohormone A neurohormone is any hormone produced and released by neuroendocrine cells (also called neurosecretory cells) into the blood. By definition of being hormones, they are secreted into the circulation for systemic effect, but they can also have a role ...
called pheromone biosynthesis activating neuropeptide (PBAN). This 33-amino-acid-long peptide is present in both sexes in the brain-
suboesophageal ganglion The suboesophageal ganglion (acronym: SOG; synonym: ''subesophageal ganglion'') of arthropods and in particular insects is part of the arthropod central nervous system (CNS). As indicated by its name, it is located ''below the'' ''oesophagus'', insi ...
s (Br-SOG) during both scotophase and photophase. It has been shown that the juvenile hormone is involved in the release of PBAN in both males and females. PBAN aids in pheromone production in females and pheromone responsiveness in males.Duportets, Line, et al. "The Pheromone Biosynthesis Activating Neuropeptide (PBAN) of the Black Cutworm Moth, ''Agrotis ipsilon'': Immunohistochemistry, Molecular Characterization and Bioassay of Its Peptide Sequence." ''Insect Biochemistry and Molecular Biology'', vol. 28, no. 8, 1998, pp. 591–599., doi:10.1016/s0965-1748(98)00033-2. In another species, PBAN release has been shown to be stimulated by external factors including photoperiod, temperature and odorants from host plantsPicimbon, Jean-Francois. "Juvenile Hormone Stimulates Pheromonotropic Brain Factor Release in the Female Black Vutworm, ''Agrotis ipsilon''". ''Journal of Insect Physiology'', vol. 41, no. 5, pp. 377–382.


Juvenile hormone

The juvenile hormone (JH), released by the
corpora allata In insect physiology and anatomy, the corpus allatum (plural: corpora allata) is an endocrine gland that generates juvenile hormone; as such, it plays a crucial role in metamorphosis. Surgical removal of the corpora allata (an allatectomy) can cau ...
(CA), is necessary for the production and release of the sex pheromone. The CA releases JH which acts on the production/release of the PBAN-like factor. So, PBAN is what connects the network in the CA to the central nervous system's production of sex pheromone. When the CA was removed, calling behavior and sex pheromone production stopped. As well, ovaries remained underdeveloped when the CA was absent. However, when decapitated females (meaning complete absence of the CA) were injected with a synthetic form of JH, ovaries were able to develop. This indicates that JH acts on the ovaries and production of sex pheromone in two independent neuroendocrine systems. In males, JH is necessary for pheromone responsiveness. When the CA was removed, males did not respond to female sex pheromones with sexual behavior. However, when the CA was implanted back, responsiveness and sexual behavior returned.


Physiology


Olfaction

''A. ipsilon'' has a sensitive
olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
system with many proteins that are expressed in the antennae. Such proteins include odorant binding proteins (OBPs), chemosensory proteins (CSPs), odorant receptors (ORs), ionotropic receptors (IRs) and sensory neuron membrane proteins (SNMPs). These proteins are responsible for recognizing sex pheromone and general odorants, such as those released by host plants.


Interactions with humans


Pest of crop plants

Each larva can consume over 400 square centimetres of foliage during its development. They feed above ground until about the fourth instar. After that they do considerable damage to crops by severing young plants at ground level. In the midwestern US, the black cutworm is considered to be a serious pest of corn. Corn is very susceptible at the one-leaf stage, but by the four- or five-leaf stage, it is relatively unaffected. Damage to the underground parts of plants can also be harmful. Other crops where serious damage occurs include cotton, maize, tobacco, sunflower, tomatoes, sugar beet and potato.


Management

There are three options to manage cutworm population and the incurred damages. Soil insecticides can be applied as a pre-plant treatment, although this may be limited by the unpredictability of cutworm population density distribution. These insecticides can also be applied as a planting-time treatment, although the same limitations still hold. The third option would be a rescue treatment that is applied after the infestations have occurred; this is also called the wait-and-see system. This may also be preferable due to a recently lower occurrence of outbreaks.Black Cutworm: Insect Fact Sheet (2003). University of Illinois. Integrated pest management, https://ipm.illinois.edu/fieldcrops/insects/black_cutworm.pdf.


See also

*
Army cutworm The army cutworm is the immature form of ''Euxoa auxiliaris''. The adult moth is called a "miller moth" because of the fine scales on its wings that rub off easily and remind people of the dusty flour that covers the clothing of a miller. These ...
(''Euxoa auxiliaris'') * Variegated cutworm (''Peridroma saucia'') * Brown cutworm (''Agrotis munda'')


References


External links

*Kimber, Ia
"73.327 BF2091 Dark Sword-grass ''Agrotis ipsilon'' (Hufnagel, 1766)"
''UKMoths''. Retrieved 28 June 2019 * Taxonomy *
''Lepiforum e.V.''
{{Taxonbar , from=Q1093329 Agrotis Moths described in 1766 Agricultural pest insects Owlet moths of Africa Cosmopolitan moths Taxa named by Johann Siegfried Hufnagel Insect pests of millets