Colorado Beetle
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The Colorado potato beetle (''Leptinotarsa decemlineata''; also known as the Colorado beetle, the ten-striped spearman, the ten-lined potato beetle, and the potato bug) is a
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
known for being a major pest of
potato The potato () is a starchy tuberous vegetable native to the Americas that is consumed as a staple food in many parts of the world. Potatoes are underground stem tubers of the plant ''Solanum tuberosum'', a perennial in the nightshade famil ...
crops. It is about long, with a bright yellow/orange body and five bold brown stripes along the length of each of its wings. Native to the
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in great-circle distance, straight-line distance from the northernmost part of Western Can ...
, it spread rapidly in potato crops across
America The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
and then
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
from 1859 onwards.
see pp. 453–454: "Doryphora, Illig.: D. 10-lineata".
/ref> The Colorado potato beetle was first observed in 1811 by
Thomas Nuttall Thomas Nuttall (5 January 1786 – 10 September 1859) was an English botanist and zoologist who lived and worked in America from 1808 until 1841. Nuttall was born in the village of Long Preston, near Settle in the West Riding of Yorkshire a ...
and was formally described in 1824 by American entomologist
Thomas Say Thomas Say (June 27, 1787 – October 10, 1834) was an American entomologist, conchologist, and Herpetology, herpetologist. His studies of insects and shells, numerous contributions to scientific journals, and scientific expeditions to Florida, Ge ...
. The beetles were collected in the
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in great-circle distance, straight-line distance from the northernmost part of Western Can ...
, where they were feeding on the buffalo bur, '' Solanum rostratum''.


Description

Adult beetles typically are in length and in width. They weigh 50–170  mg. The beetles are orange-yellow in color with 10 characteristic black stripes on their front wings or
elytra An elytron (; ; : elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometime ...
. The
specific Specific may refer to: * Specificity (disambiguation) * Specific, a cure or therapy for a specific illness Law * Specific deterrence, focussed on an individual * Specific finding, intermediate verdict used by a jury in determining the final ...
name ''decemlineata'', meaning "ten-lined", derives from this feature. Adult beetles may be visually confused with ''L. juncta'', which is the false potato beetle. Unlike the Colorado potato beetle, it is not an agricultural pest. ''L. juncta'' also has alternating black and white strips on its back, but one of the white strips in the center of each wing cover is missing and replaced by a light brown strip.


Larvae

The orange-pink larvae have a large, 9-segmented abdomen, black head, and prominent spiracles, and may measure up to in length in their final instar stage. The beetle larva has four
instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
stages. The head remains black throughout these stages, but the pronotum changes colour from black in first- and second-instar larvae to having an orange-brown edge in its third-instar. In fourth-instar larvae, about half the pronotum is coloured light brown. This tribe is characterised within the subfamily by round to oval-shaped convex bodies, which are usually brightly coloured, simple claws which separate at the base, open cavities behind the procoxae, and a variable apical segment of the maxillary palp.


Distribution

The beetle is most likely native to the area between
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
and
northern Mexico Northern Mexico ( ), commonly referred as , is an informal term for the northern cultural and geographical area in Mexico. Depending on the source, it contains some or all of the states of Baja California, Baja California Sur, Chihuahua (state), ...
, and was discovered in 1824 by Thomas Say in the Rocky Mountains. It is found in North America, and is present in every state and province except Alaska, California, Hawaii, and Nevada. It now has a wide distribution across Europe and Asia, totaling over 16 million km2. Its first association with the potato plant (''
Solanum tuberosum The potato () is a starchy tuberous vegetable native to the Americas that is consumed as a staple food in many parts of the world. Potatoes are underground stem tubers of the plant ''Solanum tuberosum'', a perennial in the nightshade famil ...
'') was not made until about 1859, when it began destroying potato crops in the region of
Omaha Omaha ( ) is the List of cities in Nebraska, most populous city in the U.S. state of Nebraska. It is located in the Midwestern United States along the Missouri River, about north of the mouth of the Platte River. The nation's List of United S ...
,
Nebraska Nebraska ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Ka ...
. Its spread eastward was rapid, at an average distance of 140 km per year. The beetle has the potential to spread to temperate areas of East Asia, India, South America, Africa, New Zealand, and Australia.


Human interaction

By 1874 it had reached the Atlantic Coast. From 1871, American entomologist
Charles Valentine Riley Charles Valentine Riley (18 September 1843 – 14 September 1895) was a British-born American entomologist and artist. He was one of the first individuals to use biological pest control and authored over 2,400 publications. He convinced Congress ...
warned Europeans about the potential for an accidental infestation caused by the transportation of the beetle from America. From 1875, several Western European countries, including Germany, Belgium, France, and Switzerland, banned imports of American potatoes to avoid infestation by ''L. decemlineata''. These controls proved ineffective, as the beetle soon reached Europe. In 1877, ''L. decemlineata'' reached the United Kingdom and was first recorded from
Liverpool Liverpool is a port City status in the United Kingdom, city and metropolitan borough in Merseyside, England. It is situated on the eastern side of the River Mersey, Mersey Estuary, near the Irish Sea, north-west of London. With a population ...
docks, but it did not become established. Many further outbreaks have occurred; the species has been eradicated in the UK at least 163 times. The last major outbreak was in 1977. It remains as a notifiable quarantine pest in the United Kingdom and is monitored by the Plant Health and Seeds Inspectorate of the Animal and Plant Health Agency (APHA) to prevent it from becoming established. A cost-benefit analysis from 1981 suggested that the cost of the measures used to exclude ''L. decemlineata'' from the UK was less than the likely costs of control if it became established. In July 2023, Colorado beetle were officially confirmed in a potato field in Kent, England. Farmers and growers, gardeners and members of the public are being encouraged to remain vigilant for signs of the pest and to report potential sightings to APHA. Elsewhere in Europe, the beetle became established near USA military bases in
Bordeaux Bordeaux ( ; ; Gascon language, Gascon ; ) is a city on the river Garonne in the Gironde Departments of France, department, southwestern France. A port city, it is the capital of the Nouvelle-Aquitaine region, as well as the Prefectures in F ...
during or immediately following
World War I World War I or the First World War (28 July 1914 – 11 November 1918), also known as the Great War, was a World war, global conflict between two coalitions: the Allies of World War I, Allies (or Entente) and the Central Powers. Fighting to ...
and had proceeded to spread by the beginning of World War II to Belgium, the Netherlands, and Spain. The population increased dramatically during and immediately following World War II and spread eastward, and the beetle is now found over much of the continent. After
World War II World War II or the Second World War (1 September 1939 – 2 September 1945) was a World war, global conflict between two coalitions: the Allies of World War II, Allies and the Axis powers. World War II by country, Nearly all of the wo ...
, in the Soviet occupation zone of Germany, almost half of all potato fields were infested by the beetle by 1950. In
East Germany East Germany, officially known as the German Democratic Republic (GDR), was a country in Central Europe from Foundation of East Germany, its formation on 7 October 1949 until German reunification, its reunification with West Germany (FRG) on ...
, they were known as ''Amikäfer'' ('Yankee beetles') following a governmental claim that the beetles were dropped by American planes. In the
European Union The European Union (EU) is a supranational union, supranational political union, political and economic union of Member state of the European Union, member states that are Geography of the European Union, located primarily in Europe. The u ...
, it remains a regulated (quarantine) pest for the Republic of Ireland, Balearic Islands, Cyprus, Malta, and southern parts of Sweden and Finland. It is not established in any of these member states, but occasional infestations can occur when, for example, wind blows adults from Russia to Finland. File:kartoffelkaefer fg01e.jpg, Native range of the potato and native and current range of the Colorado beetle File:Propagation du doryphore en Amérique du Nord.svg, Expansion of the Colorado potato beetle's range in North America, 1859–1876 File:Doryphore - expansion en Europe.svg, Expansion of the Colorado potato beetle's range in Europe, 1921–1964


Lifecycle

Colorado potato beetle females are very prolific and are capable of laying over 500 eggs in a 4- to 5-week period. The eggs are yellow to orange, and are about long. They are usually deposited in batches of about 30 eggs on the underside of host leaves. Development of all life stages depends on temperature. After 4–15 days, the eggs hatch into reddish-brown
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e with humped backs and two rows of dark brown spots, one row on each side. They feed on the leaves of their host plants. Larvae progress through four distinct growth stages (
instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
s). First instars measure about long, and the last (fourth) instars about long. The first through third instars each last about 2–3 days; the fourth lasts 4–7 days. Upon reaching full size, each fourth instar spends several days as a nonfeeding
prepupa The prepupa is a stage in the life cycle of certain Insect, insects, following the larva or Nymph (biology), nymph and preceding the pupa. It occurs in both Holometabolism, holometabolous and Hemimetabolism, hemimetabolous insects. Examples ...
, which can be recognized by its inactivity and lighter coloration. The prepupae drop to the soil and burrow to a depth of several inches, then
pupa A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
te. In 5 to 10 days, the adult beetle emerges to feed and mate. This beetle can thus go from egg to adult in as little as 21 days. Depending on temperature, light conditions, and host quality, the adults may enter
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
and delay emergence until spring. They then return to their host plants to mate and feed; overwintering adults may begin mating within 24 hours of spring emergence. In some locations, three or more generations may occur each growing season. File:Leptinotarsa-decemlineata-eggs.jpg, Eggs laid on the underside of a leaf File:Leptinotarsa decemlineata UGA1243045.jpg, 1st instar larva after hatching File:Potato beetle larvae.jpg, 3rd instar stage of larvae File:2006-07-16 larve2.jpg, 4th instar stage of larva, before pupation File:Leptinotarsa decemlineata UGA1243047 - no watermark.jpg, Pupa File:Kartoffelkäfer.jpg, Adult beetle after emergence File:Leptinotarsa decemlineata, A Coruña, Galiza.jpg, Mating adult beetles


Mate and host searching

Visual cues are important for Colorado potato beetles during the mate and host search. In a study done by Szentsi, Weber, and Jermy in the paper ''Role of visual stimuli in host and mate location of the Colorado potato beetle,'' the beetles' attraction to boards with different spectral bands, reaction to beetle-sized stationary objects, responses to such objects on boards, and attraction to prior female substances were investigated. The researchers' hypothesis was that experience with female substances would cause behavior changes in males. When shown colored boards, the beetles had a positive response between 45° and 0° in terms of mean angular directions (MADs). Beads and dead beetles without boards evoked a weaker response with MADs being variable. Colored boards and bead combinations displayed more positive MADs responses between 45° and 0°. Experience with female substances showed that male beetles showed high responses to female scent. According to the study, 43 of the 49 runs where female smear was used had a response score of 5 in contrast to the 23/42 runs without female smear receiving a score of 5. Colorado potato beetles are also attracted to the volatiles potato plants emit. In the article ''Sexual contact influences orientation to plant attractant in Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae)'' by Joseph Dickens, the beetles were attracted to
kairomone A kairomone is a semiochemical released by an organism that mediates interspecific interactions in a way that benefits a different species at the expense of the emitter. Derived from the Greek καιρός, meaning "''opportune moment""kairomon ...
substance but after mating, their attraction to it reduced. Within 24 hours of mating, there was no difference between levels of attraction to kairomone and control solvent. Lack of attraction occurred for two days but resumed three days after mating. Male beetles produce a pheromone that is further enhanced by plant host volatiles like the kairomone. After a beetle is attracted to the host, mating occurs and the female lays her eggs on the plant. The beetles' attraction to kairomone decreases until 72 hours later once oviposition occurs and the probability of re-mating increases.


Behavior and ecology


Diet

''L. decemlineata'' has a strong association with plants in the family
Solanaceae Solanaceae (), commonly known as the nightshades, is a family of flowering plants in the order Solanales. It contains approximately 2,700 species, several of which are used as agricultural crops, medicinal plants, and ornamental plants. Many me ...
, particularly those of the genus ''
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 Solana ...
''. It is directly associated with '' Solanum cornutum'' (buffalo-bur), ''
Solanum nigrum ''Solanum nigrum'', the European black nightshade or simply black nightshade or blackberry nightshade, is a species of flowering plant in the family Solanaceae, native to Eurasia and introduced in the Americas, Australasia, and South Africa. Ripe ...
'' (black nightshade), '' Solanum melongena'' (eggplant or aubergine), ''
Solanum dulcamara ''Solanum dulcamara'' is a species of vine in the genus ''Solanum'' (which also includes the potato and the tomato) of the family Solanaceae. Common names include bittersweet, bittersweet nightshade, bitter nightshade, blue bindweed, Amara Dulci ...
'' (bittersweet nightshade), '' Solanum luteum'' (hairy nightshade), ''
Solanum tuberosum The potato () is a starchy tuberous vegetable native to the Americas that is consumed as a staple food in many parts of the world. Potatoes are underground stem tubers of the plant ''Solanum tuberosum'', a perennial in the nightshade famil ...
'' (potato), and '' Solanum elaeagnifolium'' (silverleaf nightshade). They are also associated with other plants in this family, namely the species ''
Solanum lycopersicum The tomato (, ), ''Solanum lycopersicum'', is a plant whose fruit is an edible berry that is eaten as a vegetable. The tomato is a member of the nightshade family that includes tobacco, potato, and chili peppers. It originated from and was d ...
'' (tomato) and the genus ''
Capsicum ''Capsicum'' () is a genus of flowering plants in the Solanum, nightshade family Solanaceae, native to the Americas, cultivated worldwide for their edible fruit, which are generally known as "peppers" or "capsicum". Chili peppers grow on five s ...
'' (pepper).


Enemies

At least 13 insect genera, three spider
families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
, one phalangid (
Opiliones The Opiliones (formerly Phalangida) are an Order (biology), order of arachnids, Common name, colloquially known as harvestmen, harvesters, harvest spiders, or daddy longlegs (see below). , over 6,650 species of harvestmen have been discovered w ...
), and one
mite Mites are small arachnids (eight-legged arthropods) of two large orders, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as eac ...
have been recorded as either generalist or specialized predators of the varying stages of ''L. decemlineata''. These include the
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 ...
'' Lebia grandis'', the coccinellid beetles '' Coleomegilla maculata'' and '' Hippodamia convergens'', the shield bugs '' Perillus bioculatus'' and '' Podisus maculiventris'', various species of the
lacewing The Hemerobiiformia are a suborder of insects in the order Neuroptera that include most of the lacewings, antlions and their allies. The phylogeny of the Neuroptera was explored in 2014 using mitochondrial DNA sequences. The results indicate t ...
genus '' Chrysopa'', the
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 ...
genus ''
Polistes ''Polistes'' is a cosmopolitan genus of paper wasps and the only genus in the tribe Polistini. Vernacular names for the genus include umbrella wasps, coined by Walter Ebeling in 1975 to distinguish it from other types of paper wasp, in refer ...
'', and the damsel bug genus ''
Nabis Nabis may refer to: * Nabis of Sparta, reigned 207–192 BCE * Nabis (art), a Parisian post-Impressionist artistic group * ''Nabis'' (bug), a genus of insects * NABIS, National Ballistics Intelligence Service, a British government agency See a ...
''. The predatory ground beetle ''L. grandis'' is a predator of both the eggs and larvae of ''L. decemlineata'', and its larvae are
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 of the pupae. An adult ''L. grandis'' may consume up to 23 eggs or 3.3 larvae in a single day. In a laboratory experiment, ''Podisus maculiventris'' was used as a predatory threat to female ''L. decemlineata'' specimens, resulting in the production of unviable
trophic egg A trophic egg is an egg (biology), egg whose function is not reproduction but nutrition; in essence, the trophic egg serves as food for offspring hatched from viable eggs. In most species that produce them, a trophic egg is usually an Fertilization, ...
s alongside viable ones; this response to a predator ensured that additional food was available for newly hatched offspring to increase their survival rate. The same experiment also demonstrated the cannibalism of unhatched eggs by newly hatched ''L. decemlineata'' larvae as an antipredator response.


Sexual dimorphism

Colorado potato beetles exhibit
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
. In particular, they exhibit dimorphism in the adhesive tarsal setae. The paper "Sexual dimorphism in the attachment ability of the Colorado potato beetle Leptinotarsa decemlineata (Coleoptera: Chrysomelidae) to rough substrates" by Voigt demonstrates this dimorphism. The
seta In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae, ...
e, hair-like structures, in males is to help them adhere to the females' elytra when mating. Colorado potato beetles also have adhesive setae that allows them to attach to host plants. Three current setae are known: simple pointed with an asymmetric narrowing at the tip (males and females), spatula-like with a pin on its dorsal surface (males and females), setae with an adhesive terminal disc (males only). Male setae are better designed for smooth surfaces; male Colorado potato beetles have been observed attaching onto smooth glass and plastic surfaces and also attach to the smooth female elytra. Microscopy of the tarsal reveals five articulated tarsomeres and paired curved claws. Males and females have an adhesive setae covering the first three tarsomeres. The fourth is hidden and the fifth bears sensory setae with no adhesive function. Both males and females have filamentous with a tapered terminal part, lanceolate with a flattened tapered terminal part, and spatula-shaped with an enlarged tape-like terminal part. Males have a discoidal terminal part with a bulge around the disc. Female elytra appears smooth on the surface, but further magnification shows irregular lines. This indicates fluid on the elytra.


Genetics


Genetic differentiation from agriculture

Colorado potato beetles display genetic differentiation based on region. In the Columbia Basin and Central Sands, beetles in the Columbia Basin had less genetic diversity than those in Central Sands. According to the study done by Crossly, Rondon, and Schoville, in the paper ''Effects of contemporary agricultural land cover on Colorado potato beetle genetic differentiation in the Columbia Basin and Central Sands,'' nucleotide diversity in the Columbia Basin beetles ranged from 0.0056-0.0063 and 0.0073-0.0080 in Central Sands. Heterozygosity data showed the Columbia Basin was 19.4% ± 0.4% and 21.6% ± 0.8% in the Central Sands. Additional mitochondrial DNA sequencing showed two haplotypes in the Columbia Basin compared to places like Wisconsin showed seven haplotypes. Reasoning behind the genetic diversity is the landscapes of the regions: shrub-land and grains in the Columbia Basin versus the forest, corn, and beans in the Central Sands. In the same study, potatoes covered 3.5% in the Columbia Basin and 1.8% in the Central Sands. Landscape resistance can be characterized by how the land responds to the spread of beetles. Its overall effect on allele frequency covariance was low, and the Central Sands had a higher rate of decay in allele frequency. Potatoes' relative effect sizes of land cover variables on genetic differentiation was the highest in the Columbia Basin. However, when comparing all the land types, no particular land cover displayed any significant difference from the others. Genetic differentiation in the Colorado potato beetle can be impacted by agricultural practices such as
crop rotation Crop rotation is the practice of growing a series of different types of crops in the same area across a sequence of growing seasons. This practice reduces the reliance of crops on one set of nutrients, pest and weed pressure, along with the pro ...
. The same study mentioned earlier examines crop rotation's effects on genetic differentiation in Colorado potato beetles that were not found in the Central Sands. On the other hand, genetic diversity decreased with increased crop rotation in the Columbia Basin. This difference could be attributed to larger rotation differences in the Columbia Basin or differences in the landscape itself that affect the spread of the beetles. Genetic diversity is not directly impacted by the land cover type. Instead, other factors such as climate could be responsible for the differences between the Colorado potato beetle in these two regions.


Genetic differentiation due to invasion

The Colorado potato beetle has invaded North America and Europe. Because of its widespread invasion, the Colorado potato beetle displays genetic diversity in its different regions. In the paper ''The voyage of an invasive species across continents: genetic diversity of North American and European Colorado potato beetle populations'' by Grapputo, Boman, Lindström, and Mappes, sequencing of amplified mtDNA from 109 beetles in 13 populations showed 20 unique haplotypes. Three haplotypes were shared in the populations and all others were restricted to a single population in North America. 51 European beetles collected from eight populations yielded in one haplotype that was also fixed in the Idaho population. Mitochondrial data,
mtDNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the DNA contained in ...
, of North American beetles showed significant population differentiation. For example, 44% of the variation can be attributed to subdivision among populations, especially in Kentucky and Idaho. Polymorphism was highest in Colorado potato beetles in Colorado and the lowest was in France. Polymorphism and
heterozygosity Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mos ...
was higher in North America than in Europe. Heterozygosity ranged from 0.25 in New Brunswick to 0.14 in France. Further analysis revealed population differentiations between North America and Europe. There were two separate groups of European beetles, one formed by western European beetles and the second being eastern European beetles. 13% of total variation is from variation among the two continent groups, and 17% of variation is from population variance within groups. Beetles from North American and Europe formed clusters. With the exception of New Brunswick and Kentucky beetles, most beetles from the same population cluster together. In Europe, there were more complex relations between the beetles. Estonian and Spanish beetles clustered, French and Italian beetles formed separate groups, and Russian and Finnish beetles were closely related to Estonian ones. European beetles could be categorized by East and West except for Polish beetles which had relations to multiple countries.


Importance of transposable elements in genome

To help explain why Colorado Potato beetles are such difficult agricultural pests to manage and control, a group of researchers sought to test both structural and functional genetic changes in the species of beetle as compared to other arthropod species. Using community annotation, transcriptomics, and genome sequencing, they uncovered that Colorado Potato beetles have a genome consisting of several transposable elements. Transposons are sequences of genetic material that can shift/move their place within an organism's genome, and 17% of the Colorado Potato beetles’ genome consists of transposable elements. This helps explain their rapid evolution to continually resist insecticides, contributing to their global spread.


As an agricultural pest


Factors affecting beetle dispersal

Colorado potato beetles are highly mobile and are considered pests. Colorado potato beetles disperse to hosts via walking and flight. Flights have three types: short, long, and diapause. Diapause is a long-distance flight that occurs at the end of the summer. In order for dispersal to occur, certain conditions need to be met, both abiotic and biotic.


Abiotic conditions

Abiotic factors include temperature, photoperiod, insolation, wind, and gravity. A soil temperature of 9 °C causes soiled beetles to move up. They emerge when soil surface temperature is 14–15 °C. Optimal flight takeoff temperature is 27 °C. Long photoperiods enable proper flight-muscle development. Insolation is also important for flight; at least 6 hours of insolation paired with 25–28 °C temperatures are optimal for takeoff. Wind is another condition that needs to be met. Speeds of 1–3 m/s assist with takeoff for short-distance flights. Gravity can also affect flight speed in the beetles; as the Colorado potato beetle moves out of soil, it does so on slopes of 20° or more.


Biotic conditions

Biotic conditions include availability of energy reserves, insect weight, insect density, overwintered adults, and summer adults. It is speculated that proline is the primary energy substance for Colorado potato beetles in flight. Beetles that gain more than 15% of their weight after emerging fly less and for shorter distances than beetles that remain the same weight. Wing loadings for male and female beetles were 10.83 and 15.60 N/m2. Wing loading changes as beetles feed, drink, and develop eggs. Cases of large groups of beetles leaving crops have been observed when there is higher population density. However, this is likely due to destruction of the food source, not the population itself. Overwintered beetles exhibit different behavior than summer beetles. They typically fly less because it is an adaptation to the higher risks of food deprivation in the spring compared to summer. During the summer, the adults that emerge walk until they eat enough to develop proper flight muscles and develop a proper elytra.


Motivations for dispersal and stimuli

Colorado potato beetles walk in orientations to find food. In the dark, they walk at slow speeds and in circles. Beetles also move in response to olfactory cues. The beetles respond and move faster to familiar odors. Depending on satiation levels, the beetles move differently with the winds. A parallel walk with the wind is found in satiated beetles whereas starved beetles walk against it. Visual cues are also important for the beetles. Colorado potato beetles respond to light, and intensity is proportional to rest period. Beetles exhibit phototactic orientation in which they align themselves with a cone of light and move with it. Compass orientation is when large amounts of beetles walk in a single direction and have memory of their angle to the sun. The rate of linear displacement is important for the probability of the beetle finding a plant, mate, or habitat. This is important for the success of orientation mechanisms. New beetles disperse for crops once they emerge. The crops affect colonization:
crop rotation Crop rotation is the practice of growing a series of different types of crops in the same area across a sequence of growing seasons. This practice reduces the reliance of crops on one set of nutrients, pest and weed pressure, along with the pro ...
prolongs colonization, and neighboring crops are colonized rapidly and by walking. Overwintered beetles fly to find crops and once a host plant is found, flight frequency decreases. The strategy behind this is thought to be minimizing reproductive risk because female beetles that emerge in the spring are already mated. Dispersal continues after finding a host. Moving helps beetles find better resources, mates, and progeny distribution. When moving, flight is less frequent than walking in cultivated fields than in the wild. Researchers have also evaluated how flight frequency is related to the beetle's diet. In a beetle population that had returned from diapause and been exposed to poor food conditions, mean flight frequency was decreased. This is because beetles required better food conditions to regenerate their flight muscles. Prior to diapause, beetles increased their flight frequency to compensate for poor food conditions.


Potato crop pest

Around 1840, ''L. decemlineata'' adopted the cultivated potato into its host range and it rapidly became a most destructive pest of potato crops. It is today considered to be the most important insect defoliator of potatoes. It may also cause considerable damage to tomato and aubergine crops with both adults and larvae feeding on the plant's foliage. Larvae may defoliate potato plants resulting in yield losses up to 100% if the damage occurs prior to tuber formation. Larvae may consume 40 cm2 of potato leaves during the entire larval stage, but adults are capable of consuming 10 cm2 of foliage per day. The economic cost of insecticide resistance is significant, but published data on the subject are minimal. In 1994, total costs of the insecticide and crop losses in the US state of Michigan were $13.3 million, representing 13.7% of the total value of the crop. The estimate of the cost implication of insecticides and crop losses per
hectare The hectare (; SI symbol: ha) is a non-SI metric unit of area equal to a square with 100-metre sides (1 hm2), that is, square metres (), and is primarily used in the measurement of land. There are 100 hectares in one square kilometre. ...
is $138–368. Long-term increased cost to the Michigan potato industry caused by insecticide resistance in Colorado potato beetle was estimated at $0.9 to $1.4 million each year.


Potato protection

Colorado potato beetles pose significant dangers to potatoes, which are a quintessential agricultural crop. In response to the damage they do, some potatoes have been genetically modified to resist attack and damage from the beetles. Specifically, the
Russet Burbank Potato Russet Burbank is a potato cultivar with dark brown skin and few eyes that is the most widely grown potato in North America. A russet type, its flesh is white, dry, and mealy, and it is good for baking, mashing, and french fries (chips). It is a ...
. The insertion of a ''cryIIIA'' gene that codes for the insect control protein ''
Bacillus thuringiensis ''Bacillus thuringiensis'' (or Bt) is a gram-positive bacteria, 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 ...
var. Tenebrionis'' is the method that was used. Prior to its insertion, research showed that
wild-type The wild type (WT) is the phenotype of the typical form of a species as it occurs in nature. Originally, the wild type was conceptualized as a product of the standard "normal" allele at a locus, in contrast to that produced by a non-standard, " ...
''cryIIIA'' genes were expressed in low levels in plants. Plants with this gene expressed the ''cryIIIA'' protein at levels less than 0.001% of total leaf protein. Plants contain some resistance and toxicity to the Colorado potato beetles, but consistent protection requires higher levels of expression of the ''cryIIIA'' gene. Scientists modified ''cryIIIA'' by modifying the DNA protein-coding sequence without altering the
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...
sequence. The gene was transferred into the potato through a vector, specifically a ''
Agrobacterium tumefaciens ''Agrobacterium tumefaciens'' (also known as ''Rhizobium radiobacter'') is the causal agent of crown gall disease (the formation of tumours) in over 140 species of eudicots. It is a rod-shaped, Gram-negative soil bacterium. Symptoms are cause ...
'' mediated transfer. Following the introduction of the gene, Russet Burbank potato plants with the gene were tested for kanamycin resistance and Colorado potato beetle resistance. In 308 plants that were tested, 18% (55) displayed complete resistance to the beetle. Later larval stages and adult beetles are more sensitive to ''cryIIIA'' protein. Controlling adults is important because they produce the next larvae generation. Colorado potato beetles overwinter as adults in the soil and feed immediately after emerging in the spring. In ''cryIIIA'' expression levels above 0.005%, adult feeding was negligible. Oviposition was also affected. In non-
transgenic A transgene is a gene that has been transferred naturally, or by any of a number of genetic engineering techniques, from one organism to another. The introduction of a transgene, in a process known as transgenesis, has the potential to change the ...
leafs, mean number of eggs per cage were 117 and 143 in two separate trials. On the other hand, transgenic leafs displayed a mean of 1.7 and 0 eggs per cage in two trials. The female beetles were also studied, and the beetles put in the cage with transgenic plants displayed reduced size with ova that were partially or totally reabsorbed. They absorbed body fat and reproductive tissue as a result of ceasing consumption of transgenic plants. The potatoes showed benefits of the gene treatment; potatoes expressing the ''cryIIIA'' gene had protection from Colorado potato beetles in the laboratory and the field. Furthermore, these potato plants displayed agronomic and tuber characteristics that aligned with healthy Russet Burbank Potatoes.


Insecticidal management

The large-scale use of insecticides in agricultural crops effectively controlled the pest until it became resistant to
DDT Dichlorodiphenyltrichloroethane, commonly known as DDT, is a colorless, tasteless, and almost odorless crystalline chemical compound, an organochloride. Originally developed as an insecticide, it became infamous for its environmental impacts. ...
in 1952 and
dieldrin Dieldrin is an organochlorine compound originally produced in 1948 by J. Hyman & Co, Denver, as an insecticide. Dieldrin is closely related to aldrin, which reacts further to form dieldrin. Aldrin is not toxic to insects; it is oxidized in the i ...
in 1958.
Insecticide Insecticides are pesticides used to kill insects. They include ovicides and larvicides used against insect eggs and larvae, respectively. The major use of insecticides is in agriculture, but they are also used in home and garden settings, i ...
s remain the main method of pest control on commercial farms. However, many chemicals are often unsuccessful when used against this pest because of the beetle's ability to rapidly develop insecticide resistance. Different populations in different geographic regions have, between them, developed resistance to all major classes of insecticide, although not every population is resistant to every chemical. The species as a whole has evolved resistance to 56 different chemical insecticides. The mechanisms used include improved metabolism of the chemicals, reduced sensitivity of target sites, less penetration and greater excretion of the pesticides, and some changes in the behavior of the beetles. CPBs have evolved widespread insecticide resistance. No cases without fitness cost or of negative cost are known.


Nonpesticidal management

Bacterial insecticides can be effective if application is targeted towards the vulnerable early-instar larvae. Two strains of the
bacterium Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among the ...
''
Bacillus thuringiensis ''Bacillus thuringiensis'' (or Bt) is a gram-positive bacteria, 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 ...
'' produce toxins that kill the larvae. Other forms of pest control, through nonpesticidal management are available. Feeding can be inhibited by applying antifeedants, such as fungicides or products derived from Neem (''
Azadirachta indica ''Azadirachta indica'', commonly known as neem, margosa, nimtree or Indian lilac, is a tree in the mahogany family Meliaceae. It is one of the two species in the genus ''Azadirachta''. It is native to the Indian subcontinent and to parts of Sout ...
''), but these may have negative effects on the plants, as well. The steam distillate of fresh leaves and flowers of
tansy Tansy (''Tanacetum vulgare'') is a perennial, herbaceous flowering plant in the genus ''Tanacetum'' in the aster family, native to temperate Europe and Asia. It has been introduced to other parts of the world, including North America, and in ...
(''Tanacetum vulgare'') contains high levels of
camphor Camphor () is a waxy, colorless solid with a strong aroma. It is classified as a terpenoid and a cyclic ketone. It is found in the wood of the camphor laurel (''Cinnamomum camphora''), a large evergreen tree found in East Asia; and in the kapu ...
and
umbellulone Umbellulone is a headache-inducing monoterpene ketone found in the leaves of the tree '' Umbellularia californica'', sometimes known as the "headache tree". It is hypothesized to cause headaches by influencing the trigeminovascular system via TRP ...
, and these chemicals are strongly repellent to ''L. decemlineata''. ''
Beauveria bassiana ''Beauveria bassiana'' is a fungus that grows naturally in soils throughout the world and acts as a parasite on various arthropod species, causing white muscardine disease; it thus belongs to the group of entomopathogenic fungi. It is used as a ...
'' (Hyphomycetes) is a
pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
ic fungus that infects a wide range of insect species, including the Colorado potato beetle. It has shown to be particularly effective as a biological pesticide for ''L. decemlineata'' when used in combination with ''B. thuringiensis''. Crop rotation is, however, the most important cultural control of ''L. decemlineata''. Rotation may delay the infestation of potatoes and can reduce the build-up of early-season beetle populations because the adults emerging from
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
can only disperse to new food sources by walking. One 1984 study showed that rotating potatoes with nonhost plants reduced the density of early-season adults by 95.8%. Other cultural controls may be used in combination with crop rotation:
Mulch A mulch is a layer of material applied to the surface of soil. Reasons for applying mulch include conservation of soil moisture, improving soil fertility, fertility and health of the soil, reducing Weed control, weed growth, and enhancing the v ...
ing the potato crop with straw early in the growing season may reduce the beetle's ability to locate potato fields, and the mulch creates an environment that favours beetle's predators; Plastic-lined trenches have been used as pitfall traps to catch the beetles as they move toward a field of potatoes in the spring, exploiting their inability to fly immediately after emergence;
flamethrower A flamethrower is a ranged incendiary device designed to project a controllable jet of fire. First deployed by the Byzantine Empire in the 7th century AD, flamethrowers saw use in modern times during World War I, and more widely in World W ...
s may also be used to kill the beetles when they are visible at the top of the plant's foliage.


Biological management

Some potential sources of control for the Colorado golden beetle is the eulophid egg parasitoid ''Edovum puttleri''. This parasitoid can kill more than 80% of beetle eggs. It does it by parasitization, probing, and host feeding. ''Edovum'' specializes in the Colorado potato beetle, which gives it easier access to the eggs it eats. This parasitoid tolerates warmer temperatures than the beetle. The adults hunt during the warmest part of the day and have different food sources. The young feast on beetle eggs. Furthermore, these parasitoids do not overwinter which means using them for biological control requires raising them in insectaries and periodic releases. However, there are studies that attempt to genetically improve ''Edovum's'' tolerance to colder temperatures along with cultural manipulations that enable ''Edovum'' to provide useful, economic biological control. Another enemy/potential control method is a fungal pathogen called '' Beuveria bassiana''. This fungus has implications in population control of the beetles. It cannot be used to quickly contain large populations of beetles. Additionally, the pre-existing use of fungicide in disease management of crops presents an obstacle for the effectiveness of the fungus. Other reasons as to why this fungal treatment has not been utilized heavily include costs of production and longevity of formulations.


Spatial and temporal field management

Colorado Potato beetles have also shown a capacity for spatial and temporal within-field management. In one study, populations of immigrating Colorado Potato beetles were systematically targeted and their established perimeter was measured over a large field. Researchers found that immigrating adult beetles had almost no spatial dependence in any covariance-based treatment, while larval immigrant populations developed the highest densities in field centers. Results imply that perimeter tactics employed by Colorado Potato beetles can give valuable insight into site-specific resistance management programs to optimize insecticide usage. However, researchers are still not confident about the long-term effects of such resistance management programs as yield reduction requires further studying.


Relationship with humans


Cold War villain

During the
Cold War The Cold War was a period of global Geopolitics, geopolitical rivalry between the United States (US) and the Soviet Union (USSR) and their respective allies, the capitalist Western Bloc and communist Eastern Bloc, which lasted from 1947 unt ...
, some countries in the
Warsaw Pact The Warsaw Pact (WP), formally the Treaty of Friendship, Co-operation and Mutual Assistance (TFCMA), was a Collective security#Collective defense, collective defense treaty signed in Warsaw, Polish People's Republic, Poland, between the Sovi ...
claimed that the beetles had been introduced by the
CIA The Central Intelligence Agency (CIA; ) is a civilian foreign intelligence service of the federal government of the United States tasked with advancing national security through collecting and analyzing intelligence from around the world and ...
in an attempt to reduce food security by destroying the agriculture of the
Soviet Union The Union of Soviet Socialist Republics. (USSR), commonly known as the Soviet Union, was a List of former transcontinental countries#Since 1700, transcontinental country that spanned much of Eurasia from 1922 until Dissolution of the Soviet ...
.Ukraine's Reins Weaken as Chaos Spreads
The New York Times ''The New York Times'' (''NYT'') is an American daily newspaper based in New York City. ''The New York Times'' covers domestic, national, and international news, and publishes opinion pieces, investigative reports, and reviews. As one of ...
(4 May 2014)Lyashko in Lviv poured green
Ukrayinska Pravda ''Ukrainska Pravda'' is a Ukrainian socio-political online media outlet founded by Heorhii Gongadze in April 2000. After Gongadze’s death in September 2000, the editorial team was led by co-founder Olena Prytula, who remained the editor-in ...
(18 June 2014)
A widespread campaign was launched against the beetles; posters were put up and school children were mobilized to gather the pests and kill them in
benzene Benzene is an Organic compound, organic chemical compound with the Chemical formula#Molecular formula, molecular formula C6H6. The benzene molecule is composed of six carbon atoms joined in a planar hexagonal Ring (chemistry), ring with one hyd ...
or spirit.


Philately

''L. decemlineata'' is an iconic species and has been used as an image on stamps because of its association with the recent history of both North America and Europe. For example, in 1956, Romania issued a set of four stamps calling attention to the campaign against insect pests, and it was featured on a 1967 stamp issued in
Austria Austria, formally the Republic of Austria, is a landlocked country in Central Europe, lying in the Eastern Alps. It is a federation of nine Federal states of Austria, states, of which the capital Vienna is the List of largest cities in Aust ...
. The beetle also appeared on stamps issued in Benin, Tanzania, the United Arab Emirates, and Mozambique.


In popular culture

Neapolitan
mandolin A mandolin (, ; literally "small mandola") is a Chordophone, stringed musical instrument in the lute family and is generally Plucked string instrument, plucked with a plectrum, pick. It most commonly has four Course (music), courses of doubled St ...
s (also called Italian mandolins) are often called ''tater bugs'', a nickname given by American
luthier A luthier ( ; ) is a craftsperson who builds or repairs string instruments. Etymology The word ' is originally French and comes from ''luth'', the French word for "lute". The term was originally used for makers of lutes, but it came to be ...
Orville Gibson Orville H. Gibson (May 8, 1856 – August 19, 1918) was an American luthier who founded the Gibson (guitar company), Gibson Guitar Company in Kalamazoo, Michigan in 1902, makers of guitars, mandolins and other instruments. His earliest known ins ...
, because the shape and stripes of the different color wood strips resemble the back of the Colorado beetle. The fans of
Alemannia Aachen Aachener Turn- und Sportverein Alemannia 1900 e. V., short Alemannia Aachen (), is a German football club from the western city of Aachen, North Rhine-Westphalia. A long-term fixture of the country's second division, Alemannia enjoyed a three- ...
carry the nickname "Kartoffelkäfer", from the German name for the Colorado beetle, because of striped yellow-black jerseys of the team. During the
2014 pro-Russian unrest in Ukraine From the end of February 2014, in the aftermath of the Euromaidan and the Revolution of Dignity, which resulted in the ousting of Russian-leaning President of Ukraine, Ukrainian President Viktor Yanukovych, demonstrations by Russian-backed, ...
, the word , from the Ukrainian and Russian term for Colorado beetle (, ), gained popularity among Ukrainians as a derogatory term to describe pro-Russian separatists in the
Donetsk Donetsk ( , ; ; ), formerly known as Aleksandrovka, Yuzivka (or Hughesovka), Stalin, and Stalino, is an industrial city in eastern Ukraine located on the Kalmius River in Donetsk Oblast, which is currently occupied by Russia as the capita ...
and
Luhansk Luhansk (, ; , ), also known as Lugansk (, ; , ), is a city in the Donbas in eastern Ukraine. As of 2022, the population was estimated to be making Luhansk the Cities in Ukraine, 12th-largest city in Ukraine. Luhansk served as the administra ...
Oblasts (provinces) of
Eastern Ukraine Eastern Ukraine or East Ukraine (; ) is primarily the territory of Ukraine east of the Dnipro (or Dnieper) river, particularly Kharkiv, Luhansk and Donetsk oblasts (provinces). Dnipropetrovsk and Zaporizhzhia oblasts are often also regarded as ...
. The nickname reflects the similarity of black and orange stripes on St. George's ribbons worn by many of the separatists. In some European cultures, the Colorado potato beetle is known as the '
gourd Gourds include the fruits of some flowering plant species in the family Cucurbitaceae, particularly '' Cucurbita'' and '' Lagenaria''. The term refers to a number of species and subspecies, many with hard shells, and some without. Many gourds ha ...
beetle' due to the likeness of the beetle to various gourds of the
Cucurbitaceae The Cucurbitaceae (), also called cucurbits or the gourd family, are a plant family (biology), family consisting of about 965 species in 101 genera.
family.


Notes

:1.For a more comprehensive list of natural predators, pathogens and parasitoids, se
here


References


External links


PotatoBeetle.orgPolish Film Chronicle: Fight against Colorado Beetle
{{Authority control Chrysomelinae Potatoes Agricultural pest insects Beetles of the United States Fauna of the Western United States Beetles described in 1824 Articles containing video clips Taxa named by Thomas Say Insects in culture Beetles of Europe Biological anti-agriculture weapons