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American bullfrog

The American bullfrog (Lithobates catesbeianus), often simply known as the bullfrog in Canada and the United States, is a large true frog native to eastern North America. It typically inhabits large permanent water bodies such as swamps, ponds, and lakes. Bullfrogs can also be found in manmade habitats such as pools, koi ponds, canals, ditches and culverts. The bullfrog gets its name from the sound the male makes during the breeding season, which sounds similar to a bull bellowing. The bullfrog is large and is commonly eaten throughout its range, especially in the southern United States where they are plentiful.

Their presence as a food source has led to bullfrogs being distributed around the world outside of their native range. Bullfrogs have been introduced into the Western United States, South America, Western Europe, China, Japan, and southeast Asia. In these places they are invasive species due to their voracious appetite and the large number of eggs they produce, which has a negative effect on native amphibians and other fauna. Bullfrogs are very skittish which makes capture difficult and so they often become established.

Other than for food, bullfrogs are also used for dissection in science classes. Albino bullfrogs are sometimes kept as pets, and bullfrog tadpoles are often sold at ponds or fish stores.

Taxonomy
Some authorities use the scientific name, Lithobates catesbeiana, although others prefer Rana catesbeiana.

Genome
The nuclear genome (~5.8Gbp) of the North American bullfrog (Rana [Lithobates] catesbeiana) was published in 2017 and provides a resource for future Ranidae research.

Etymology
The specific name, catesbeiana (feminine) or catesbeianus (masculine), is in honor of English naturalist Mark Catesby.

Description
The dorsal (upper) surface of the bullfrog has an olive-green basal color, either plain or with mottling and banding of grayish brown. The ventral (under) surface is off-white blotched with yellow or gray. Often, a marked contrast in color is seen between the green upper lip and the pale lower lip. The teeth are tiny and are useful only in grasping. The eyes are prominent with brown irises and horizontal, almond-shaped pupils. The tympana (eardrums) are easily seen just behind the eyes and the dorsolateral folds of skin enclose them. The limbs are blotched or banded with gray. The fore legs are short and sturdy and the hind legs are long. The front toes are not webbed, but the back toes have webbing between the digits with the exception of the fourth toe, which is unwebbed.

Bullfrogs are sexually dimorphic, with males being smaller than females and having yellow throats. Males have tympana larger than their eyes, whereas the tympana in females is about the same size as the eyes. Bullfrogs measure about 3.6 to 6 in in snout–to–vent length. They grow fast in the first eight months of life, typically increasing in weight from 5 to 175 g, and large, mature individuals can weigh up to 500 g. In some cases bullfrogs have been recorded as attaining 800 g and measuring up to 8 in from snout to vent. The American bullfrog is the largest species of true frog in North America.

Online Model Organism Database
xenbase provides limited support (BLAST, JBrowse tracks, genome download) for the bullfrog.

Distribution
The bullfrog is originally native to eastern North America, where it is commonly found in every U.S. state east of the Mississippi River. Its natural range extends from the eastern Canadian Maritime Provinces to as far west as Idaho and Texas, and as far north as Michigan (including the Upper Peninsula), Minnesota and Montana; it is largely absent in North Dakota. The bullfrog has also been introduced onto Nantucket island, as well as portions of the western U.S., including Arizona, Colorado, Hawaii, Idaho, Nevada, New Mexico, Oregon, Utah, Washington and Wyoming. In these states, it is considered to be an invasive species, as concerns exist that it may outcompete or prey upon native species of reptiles and amphibians, disrupting the delicate ecological balance of certain areas. The bullfrog has been introduced in Hawaii, South America, Asia, the Caribbean, and Europe for various purposes including frog farming and population control of other species. It is very common on the West Coast, especially in California, where it is believed to pose a threat to the California red-legged frog, and is considered to be a factor in the decline of that vulnerable species. Bullfrogs have been found to feed on the young of several snakes, including the California endemic giant garter snake, a threatened species. In early 2023, the Utah Department of Natural Resources began tweeting tips on how to catch and cook bullfrogs in an effort to encourage residents to help control the growing population by catching the invasive frogs for food.

Other countries and regions into which the bullfrog has been introduced include the extreme south of British Columbia, Canada, nearly every state in Mexico, as well as Belgium, Cuba, the Dominican Republic, Haiti, Italy, Jamaica, the Netherlands, and Puerto Rico. It is also found in Argentina, Brazil, China, Colombia, Japan, South Korea, Uruguay and Venezuela. The reasons for introducing the bullfrog to these areas have largely been intentional, either to provide humans with a source of food or as biological control agents. In addition to the unintended escape of frogs from breeding establishments or scientific research facilities, captive escapees or released  pets are also a possibility. Conservationists are concerned that the bullfrog is relatively immune to the fungal infection chytridiomycosis (also called ‘chytrid’ fungus) which has been ravaging numerous frog species, and, as it invades new territories, it may assist in the spread of this lethal fungus as an asymptomatic carrier to the more susceptible, native species of frog it encounters.

Breeding behavior
The bullfrog breeding season typically lasts two to three months. A study of bullfrogs in Michigan showed the males arriving at the breeding site in late May or early June, and remaining in the area into July. The territorial males that occupy sites are usually spaced some 3 to 6 m apart and call loudly. At least three different types of calls have been noted in male bullfrogs under different circumstances. These distinctive calls include territorial calls made as threats to other males, advertisement calls made to attract females, and encounter calls which precede combat.

The bullfrogs have a prolonged breeding season, with the males continuously engaging in sexual activity throughout. Males are present at the breeding pond for longer periods than females during the entire season, increasing their chances of multiple matings. The sex ratio is typically skewed toward males. Conversely, females have brief periods of sexual receptivity during the season. In one study, female sexual activity typically lasted for a single night and mating did not occur unless the females initiated the physical contact. Males only clasp females after they have indicated their willingness to mate. This finding refutes previous claims that a male frog will clasp any proximate female with no regard to whether the female has consented. Once a male finds a receptive female he will clasp onto her and undergo amplexus—reproductive position—by utilization of the males' forelimbs. The enlargement of forelimb muscles is a sexually dimorphic trait seen in the male bullfrog. One study investigating male and female bullfrog forelimbs muscles found males had significantly stronger muscles that could undergo longer durations of activity before the onset of fatigue. The significance of forelimb sexual dimorphism allow males to remain in amplexus with the female for longer durations increasing their chance at reproductive success in the highly competitive mating environment.

These male and female behaviors cause male-to-male competition to be high within the bullfrog population and sexual selection for the females to be an intense process. Kentwood Wells postulated leks, territorial polygyny, and harems are the most likely classifications for the bullfrog mating system. Leks would be a valid description because males congregate to attract females, and the females arrive to the site for the purpose of copulation. In a 1980 study on bullfrogs in New Jersey, the mating system was classified as resource-defense polygyny. The males defended territories within the group and demonstrated typical physical forms of defense.

Choruses
Male bullfrogs aggregate into groups called choruses. The male chorus behavior is analogous to the lek formation of birds, mammals, and other vertebrates. Choruses are dynamic, forming and remaining associated for a few days, breaking down temporarily, and then forming again in a new area with a different group of males. Male movement has experimentally been noted to be dynamic. In the Michigan study, the choruses were described as "centers of attraction" in which their larger numbers enhanced the males' overall acoustical displays. This is more attractive to females and also attractive to other sexually active males. Choruses in this study were dynamic, constantly forming and breaking up. New choruses were formed in other areas of the site. Males moved around and were highly mobile within the choruses.

A review of multiple studies on bullfrogs and other anurans noted male behavior within the groups changes according to the population density of the leks. At higher population densities, leks are favored due to the difficulty in defending individual territories among a large population of males. This variance causes differences in how females choose their mates. When the male population density is low and males maintain clearer, more distinct territories, female choice is mostly determined by territory quality. When male population density is higher, females depend on other cues to select their mates. These cues include the males' positions within the chorus and differences in male display behaviors among other determinants. Social dominance within the choruses is established through challenges, threats, and other physical displays. Older males tend to acquire more central locations while younger males were restricted to the periphery.

Chorus tenure is the number of nights that a male participates in the breeding chorus. One study distinguishes between chorus tenure and dominant tenure. Dominant tenure is more strictly defined as the amount of time a male maintains a dominant status. Chorus tenure is restricted due to increased risk of predation, lost foraging opportunities, and higher energy consumption. Calling is postulated to be energetically costly to anurans in general. Energy is also expended through locomotion and aggressive interactions of male bullfrogs within the chorus.

Aggressive behavior
To establish social dominance within choruses, bullfrogs demonstrate various forms of aggression, especially through visual displays. Posture is a key factor in establishing social position and threatening challengers. Territorial males have inflated postures while non-territorial males remain in the water with only their heads showing. For dominant (territorial) males, their elevated posture reveals their yellow-colored throats. When two dominant males encounter each other, they engage in a wrestling bout. The males have their venters clasped, each individual in an erect position rising to well above water level. The New Jersey study noted the males would approach each other to within a few centimeters and then tilt back their heads, displaying their brilliantly colored gular sacs. The gular is dichromatic in bullfrogs, with dominant and fitter males displaying yellow gulars. The New Jersey study also reported low posture with only the head exposed above the water surface was typical of subordinate, or non-territorial males, and females. High posture was demonstrated by territorial males, which floated on the surface of the water with their lungs inflated, displaying their yellow gulars. Males optimize their reproductive fitness in a number of ways. Early arrival at the breeding site, prolonged breeding with continuous sexual activity throughout the season, ownership of a centrally located territory within the chorus, and successful movement between the dynamically changing choruses are all common ways for males to maintain dominant, or territorial, status within the chorus. Older males have greater success in all of these areas than younger males. Some of the males display a more inferior role, termed by many researchers as the silent male status. These silent males adopt a submissive posture, sit near resident males and make no attempt to displace them. The silent males do not attempt to intercept females but are waiting for the territories to become vacant. This has also been called the alternate or satellite male strategy.

Growth and development
After selecting a male, the female deposits eggs in his territory. During the mating grasp, or amplexus, the male rides on top of the female, grasping her just behind her fore limbs. The female chooses a site in shallow water among vegetation, and lays a batch of up to 20,000 eggs, and the male simultaneously releases sperm, resulting in external fertilization. The eggs form a thin, floating sheet which may cover an area of 0.5 to 1.0 m2. The embryos develop best at water temperatures between 24 and 30 °C and hatch in three to five days. If the water temperature rises above 32 °C, developmental abnormalities occur, and if it falls below 15 °C, normal development ceases. Newly hatched tadpoles show a preference for living in shallow water on fine gravel bottoms. American bullfrog tadpoles have also "showed a preference for habitats containing structure." This may reflect a lesser number of predators in these locations. As they grow, they tend to move into deeper water. The tadpoles initially have three pairs of external gills and several rows of labial teeth. They pump water through their gills by movements of the floor of their mouths, trapping bacteria, single-celled algae, protozoans, pollen grains, and other small particles on mucus in a filtration organ in their pharynges. As they grow, they begin to ingest larger particles and use their teeth for rasping. They have downward-facing mouths, deep bodies, and tails with broad dorsal and ventral fins. Time to metamorphosis ranges from a few months in the southern part of the range to 3 years in the north, where the colder water slows development. Maximum lifespan in the wild is estimated to be 8 to 10 years, but one frog lived for almost 16 years in captivity.

Feeding
Bullfrogs are voracious, opportunistic, ambush predators that prey on any small animal they can overpower and consume. Bullfrog stomachs have been found to contain rodents, small lizards and snakes, other frogs and toads, other amphibians, crayfish, other crustaceans, small birds, scorpions, tarantulas and bats, as well as the many types of invertebrates, such as snails, worms and insects, which are the usual food of ranid frogs. These studies revealed the bullfrog's diet to be unique among North American ranids in the inclusion of a large percentage of aquatic animals, such as fish, tadpoles, ram's horn snails, and dytiscid beetles, as well as the aquatic eggs of fish, frogs, insects, or salamanders. Cannibalism has been observed in bullfrog populations in resource-limited environments. Bullfrogs are able to capture large, strong prey because of the powerful grip of their jaws after the initial ranid tongue strike. However, there is a correlation found with size of prey relative to body size of the bullfrog. Juveniles and adults typically go after prey that is relative to their own body size. The bullfrog is able to make allowance for light refraction at the water-air interface by striking at a position posterior to the target's perceived location. The comparative ability of bullfrogs to capture submerged prey, compared to that of the green frog, leopard frog, and wood frog (L. clamitans, L. pipiens, and L. sylvaticus, respectively) was also demonstrated in laboratory experiments.

Prey motion elicits feeding behavior. First, if necessary, the frog performs a single, orienting bodily rotation ending with the frog aimed towards the prey, followed by approaching leaps, if necessary. Once within striking distance, the bullfrog begins its feeding strike, which consists of a ballistic lunge (eyes closed as during all leaps) that ends with the mouth opening. At this stage, the fleshy, mucus-coated tongue is extended towards the prey, often engulfing it, while the jaws continue their forward travel to close (bite) just as the tongue is retracted. Large prey that do not fit entirely into the mouth are stuffed in with the hands. In laboratory observations, bullfrogs taking mice usually swam underwater with prey in mouth, apparently with the advantageous result of altering the mouse's defense from counter-attack to struggling for air. Asphyxiation is the most likely cause of death of warm-blooded prey.

Biomechanical background of tongue projection
The speed of a bullfrog's tongue strike is much faster than it should be if muscles were the only force behind it. Similar to the tension on a slingshot pulled all the way back, when the frog's mouth is closed, tension is put into the elastic tissues of the tongue, and also into the elastic tendons of the lower jaw. When the frog attacks prey, opening its mouth is like letting go of the slingshot; the elastic force stored up in both the tongue and the jaw are combined to shoot the tip of the tongue toward the prey much faster than the prey's ability to see the strike and evade capture, completing the strike and retrieval in approximately 0.07 seconds. Another benefit of this elastic-force based attack is that it is not dependent on background temperature. A frog with a cold body temperature has muscles that move more slowly, but it can still attack prey with the same speed as if its body was warm.

Ballistic tongue projection of the related leopard frog is possible due to the presence of elastic structures that allow storage and subsequent release of elastic recoil energy. This accounts for the tongue projecting with higher power output than would develop by muscular action alone. Also, such mechanism relieves the tongue's musculature from physiological constraints such as limited peak power output - mechanical efficiency and thermal dependence by uncoupling the activation of the depressor mandibulae's contractile units from actual muscular movement. In other words, the kinematic parameters developed by contribution of the elastic structures differ from those developed by muscular projection, accounting for the difference in velocity, power output, and thermal dependence.

Ecology
Bullfrogs are an important item of prey to many birds (especially large herons), North American river otters (Lontra canadensis), predatory fish, and occasionally other amphibians. Predators of American bullfrogs once in their adult stages can range from 150 g belted kingfishers (Megaceryle alcyon) to 1100 lb American alligators (Alligator mississippiensis). The eggs and larvae are unpalatable to many salamanders and fish, but the high levels of activity of the tadpoles may make them more noticeable to a predator not deterred by their unpleasant taste. Humans hunt bullfrogs as game and consume their legs. Adult frogs try to escape by splashing and leaping into deep water. A trapped individual may squawk or emit a piercing scream, which may surprise the attacker sufficiently for the frog to escape. An attack on one bullfrog is likely to alert others in the vicinity to danger and they will all retreat into the safety of deeper water. Bullfrogs may be at least partially resistant to the venom of copperhead (Agkistrodon contortrix) and cottonmouth (Agkistrodon piscivorus) snakes, though these species are known natural predators of bullfrogs as are northern water snakes (Nerodia sipedon).

Considering the invasive nature of the L. catesbeianus, multiple traits within the species contribute to its competitive ability. The generalist diet of the American bullfrog allows for it to consume food in different environments. When observing the contents of American bullfrog stomachs, it was discovered that adult bullfrogs regularly consume predators of bullfrog young, including dragonfly nymphs, garter snakes, and giant water bugs. Thus, the ecological check on American bullfrog juveniles in invaded areas become less effective. L. catesbeianus seems to exhibit traits of immunity or resistance against the antipredator defenses of other organisms. Analysis of stomach contents from bullfrog populations in New Mexico show the regular consumption of wasps, with no conditioned avoidance due to the wasps' stingers. Along the Colorado river, L. catesbeianus stomach contents indicate the ability to withstand the discomforting spines of the stickleback fish. Reports of American bullfrogs eating scorpions and rattlesnakes also exist.

Analysis of the American bullfrog's realized niche at various sites in Mexico, and comparisons with the niches of endemic frogs show that it is possible that the American bullfrog capable of niche shift, and pose a threat to many endemic Mexican frog species, even those that are not currently in competition with the American bullfrog.

Invasive species
In areas where the American bullfrog is introduced, the population can be controlled by various means. One project (3n-Bullfrog project) uses sterile triploïd (3n) bullfrogs. In Europe, the American bullfrog is included since 2016 in the list of Invasive Alien Species of Union concern (the Union list). This implies that this species cannot under any circumstances be imported, bred, transported, commercialized, or intentionally released into the environment in the whole of the European Union. The American bullfrog has been known to spread the amphibian pathogen Batrachochytrium dendrobatidis among populations that is has been introduced to.

Human use
The American bullfrog provides a food source, especially in the Southern and some areas of the Midwestern United States. The traditional way of hunting them is to paddle or pole silently by canoe or flatboat in ponds or swamps at night; when the frog's call is heard, a light is shone at the frog which temporarily inhibits its movement. The frog will not jump into deeper water as long as it is approached slowly and steadily. When close enough, the frog is gigged with a multiple-tined spear and brought into the boat. Bullfrogs can also be stalked on land, by again taking great care not to startle them. In some states, breaking the skin while catching them is illegal, and either grasping gigs or hand captures used. The only parts normally eaten are the rear legs, which resemble small chicken drumsticks, have a similar flavor and texture and can be cooked in similar ways.

Commercial bullfrog culture in near-natural enclosed ponds has been attempted, but is fraught with difficulties. Although pelleted feed is available, the frogs will not willingly consume artificial diets, and providing sufficient live prey is challenging. Disease also tends to be a problem even when great care is taken to provide sanitary conditions. Other challenges to be overcome may be predation, cannibalism, and low water quality. The frogs are large, have powerful leaps, and inevitably escape after which they may wreak havoc among the native frog population. Countries that export bullfrog legs include the Netherlands, Belgium, Mexico, Bangladesh, Japan, China, Taiwan, and Indonesia. Most of these frogs are caught from the wild, but some are captive-reared. The United States is a net importer of frog legs.

The American bullfrog is used as a specimen for dissection in many schools across the world. It is the state amphibian of Missouri, Ohio, and Oklahoma.

title: American bullfrog


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Haapsalu

Haapsalu is a seaside resort town located on the west coast of Estonia. It is the administrative centre of Lääne County, and on 1 January 2020 it had a population of 9,375.

History
The name "Haapsalu" derives from Estonian words haab 'aspen' and salu (archaic meaning) 'island'. Until the first half of the 20th century, in Swedish and German the town was called Hapsal. Haapsalu and the surrounding area was the center for the Estonian Swedes from the 13th century until the evacuation of almost all ethnic Swedes from Estonia prior to the Soviet invasion of Estonia during World War II in 1944.

The first written record of Hapsal dates back to 1279, when the town was chartered and became the capital of the Bishopric of Ösel-Wiek, which it remained for the following three centuries. Buildings from those early days remain today, including an episcopal castle which has the largest single-nave church in Estonia.

Neighborhoods of Haapsalu
There are six neighborhoods of Haapsalu:
 * Holmi
 * Kesklinn
 * Männiku
 * Paralepa
 * Randsalu
 * Vanalinn.

Healing by sea mud
The sea mud in Haapsalu is claimed to have a curative effect. A military doctor, Carl Abraham Hunnius, founded the first mud cure resort in 1825. News of the "curative mud" spread quickly amongst the wealthy customers in then capital city Saint Petersburg, and elsewhere in the former Russian Empire. The mud spas were frequented by the Russian imperial Romanov family. For almost 200 years, Haapsalu has been a popular summer destination where people from all around the world come for medical treatment. At present, there are three "mud cure" establishments in Haapsalu.

Other attractions
The Land of Ilon Wikland (Wiklandia), a recreation centre for children, is set to open in a few years within the town. Wikland, a famous book illustrator, has had a strong bond with Haapsalu since her childhood.

The August Blues Festival is held every year in August in Haapsalu.

Since 2005, the town hosts Haapsalu Horror and Fantasy Film Festival, an annual film festival dedicated to genre films. In 2017, the pastors of Haapsalu made an open statement calling to end the city's financing of the festival, claiming the horror and violence depicted in the screened films were not fit to represent the resort town image. The same year the festival was held to a record-breaking attendance.

In popular culture
Pyotr Ilyich Tchaikovsky in 1867 wrote a suite of three pieces for piano during his stay in Haapsalu, titled Souvenir de Hapsal.

In the 19th century, the town became famous for its "Haapsalu shawls", a delicate craft made by local women.

Haapsalu has sometimes been called the "Venice of the Baltics" — an apparent exaggeration used mostly to promote the resort town to foreign tourists.

Haapsalu is site of a fencing school founded by Estonian fencer Endel Nelis, used as the setting of the Finnish-Estonian film The Fencer.

license: Creative Commons - Attribution Share-Alike - https://creativecommons.org/licenses/by-sa/4.0/

title: Haapsalu


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User talk:Reihaneh

Hello new user and welcome to Wikipedia. A few tips for you:

 * Peruse Welcome, newcomers and associated pages
 * Sign talk page entries with ~, which is automatically converted to a name and date
 * Ask questions on Village pump
 * Have fun.

-- Alexandros 20:28, 25 Nov 2003 (UTC)

title: User talk:Reihaneh


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User:Ooterness

I'll just put some quotes here until I think of something better:

"The checks and balances of democratic governments were invented because human beings realized how unfit they were to govern themselves. They needed a system, yes, an industrial-age machine" -Helios, a intelligent supercomputer from Deus_Ex

I think Deus_Ex is quite possibly the best game ever, partly because it is filled with political commentary. In terms of this particular quote, I agree that the system-like nature of such constructions (and the timing of their widespread emergence) are probably not a coincidence in terms of the revolutionary new ways of looking at the world and how it ought to function. That an AI would seize upon this idea is, of course, a bit narcisistic, but that's Helios. I'm definitely looking forward to Deus Ex 2, which supposedly will allow players to choose nearly any allegiance based on their preferred ideologies.

"I'll bet ye've got all *sorts* o' barmy questions! 'Can you tell me about this place?  Who's the Lady o' Pain?  I'm lookin' fer the magic Girdle o' Swank Iron, have ye seen it?  Do ye know where the Holy Flamin' Frost-Brand Gronk-Slayin' Vorpal Hammer o' Woundin' an' Returnin' an' Shootin'-Lightnin'-Out-Yer-Bum is?'" -Old lady, Planescape_Torment

It's funny because it's true; most "characters" in RPGs are only looking for items and XP.

title: User:Ooterness


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Valentinian

Valentinian may refer to:
 * Valentinian I or Valentinian the Great (321–375), Western Roman emperor from 364 to 375
 * Valentinian II (371–392), Western Roman Emperor from 375 to 392
 * Valentinian III (419–455), Western Roman Emperor from 425 to 455
 * Valentinus (Gnostic), theologian and founder of Valentinianism
 * Valentinian (play), a Jacobean-era stage play
 * Valentinianic dynasty, an Imperial Roman dynasty founded by Valentinian I and sometimes known as the Valentinian dynasty

title: Valentinian


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Colorado potato beetle

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 known for being a major pest of potato crops. It is about 10 mm long, with a bright yellow/orange body and five bold brown stripes along the length of each of its elytra. Native to the Rocky Mountains, it spread rapidly in potato crops across America and then Europe from 1859 onwards.

The Colorado potato beetle was first observed in 1811 by Thomas Nuttall and was formally described in 1824 by American entomologist Thomas Say. The beetles were collected in the Rocky Mountains, where they were feeding on the buffalo bur, Solanum rostratum.

Description
Adult beetles typically are 6 - 11 mm in length and 3 mm 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. The specific 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 15 mm in length in their final instar stage.

The beetle larva has four instar 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 and northern Mexico, 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) was not made until about 1859, when it began destroying potato crops in the region of Omaha, Nebraska. 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 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 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 1976. It remains as a notifiable quarantine pest in the United Kingdom and is monitored by DEFRA 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.

Elsewhere in Europe, the beetle became established near USA military bases in Bordeaux during or immediately following World War I 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, in the Soviet occupation zone of Germany, almost half of all potato fields were infested by the beetle by 1950. In East Germany, they were known as Amikäfer ('Yankee beetles') following a governmental claim that the beetles were dropped by American planes. In the European Union, 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.

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 1 mm 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 larvae 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 (instars). First instars measure about 1.50 mm long, and the last (fourth) instars about 8 mm 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, 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 pupate. 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 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.

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 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.

Diet
L. decemlineata has a strong association with plants in the family Solanaceae, particularly those of the genus Solanum. It is directly associated with Solanum cornutum (buffalo-bur), Solanum nigrum (black nightshade), Solanum melongena (eggplant or aubergine), Solanum dulcamara (bittersweet nightshade), Solanum luteum (hairy nightshade), Solanum tuberosum (potato), and Solanum elaeagnifolium (silverleaf nightshade). They are also associated with other plants in this family, namely the species Solanum lycopersicum (tomato) and the genus Capsicum (pepper).

Enemies

At least 13 insect genera, three spider families, one phalangid (Opiliones), and one mite have been recorded as either generalist or specialized predators of the varying stages of L. decemlineata. These include the ground beetle Lebia grandis, the coccinellid beetles Coleomegilla maculata and Hippodamia convergens, the shield bugs Perillus bioculatus and Podisus maculiventris, various species of the lacewing genus Chrysopa, the wasp genus Polistes, and the damsel bug genus Nabis.

The predatory ground beetle L. grandis is a predator of both the eggs and larvae of L. decemlineata, and its larvae are parasitoids 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 eggs 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. 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 setae, 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.

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. 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, 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 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.

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 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 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. The insertion of a cryIIIA gene that codes for the insect control protein ''Bacillus thuringiensis var. Tenebrionis'' is the method that was used. Prior to its insertion, research showed that wild-type 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 sequence. The gene was transferred into the potato through a vector, specifically a Agrobacterium tumefaciens 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 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 in 1952 and dieldrin in 1958. Insecticides 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 Bacillus thuringiensis 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), but these may have negative effects on the plants, as well. The steam distillate of fresh leaves and flowers of tansy (Tanacetum vulgare) contains high levels of camphor and umbellulone, and these chemicals are strongly repellent to L. decemlineata.

Beauveria bassiana (Hyphomycetes) is a pathogenic 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 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: Mulching 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; flamethrowers 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 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.

Cold War villain
During the Cold War, some countries in the Warsaw Pact claimed that the beetles had been introduced by the CIA in an attempt to reduce food security by destroying the agriculture of the Soviet Union. 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 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. The beetle also appeared on stamps issued in Benin, Tanzania, the United Arab Emirates, and Mozambique.

In popular culture
Neapolitan mandolins (also called Italian mandolins) are often called tater bugs, a nickname given by American luthier Orville Gibson, because the shape and stripes of the different color wood strips resemble the back of the Colorado beetle.

The fans of Alemannia Aachen 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, the word, from the Ukrainian and Russian term for Colorado beetle (жук колорадський, колорадский жук), gained popularity among Ukrainians as a derogatory term to describe pro-Russian separatists in the Donetsk and Luhansk Oblasts (provinces) of Eastern Ukraine. The nickname reflects the similarity of black and orange stripes on St. George's ribbons worn by many of the separatists.

title: Colorado potato beetle


text:
Eric Vloeimans

Eric Vloeimans (born 24 March 1963) is a Dutch musician, songwriter, and record producer.

Biography

Although he studied classical music as a child, he became interested in jazz at the Rotterdam Academy of Music. After graduating in 1988, he moved to New York City and studied trumpet with Donald Byrd. He was a member of big bands led by Mercer Ellington and Frank Foster. In the 1990s, he recorded the album Bitches and Fairy Tales (1999) with Marc Johnson, Joey Baron, and John Taylor. Vloeimans has also worked with Kinan Azmeh, Michiel Borstlap, Lars Danielsson, Jimmy Haslip, Joe LaBarbera, Nguyên Lê, Ernst Reijseger, and Bugge Wesseltoft.

Awards and honors

 * Edison Award, Bitches and Fairy Tales, 1999
 * Boy Edgar Award, Umai, 2001
 * Bird Award, Best Jazz Trumpeter, 2002
 * Gouden Nutcracker, Dutch Jazz Album, Heavensabove, 2011

As leader

 * No Realistics (Art in Jazz, 1992)
 * First Floor (Challenge, 1994)
 * Bestiarium (Challenge, 1996)
 * Bitches and Fairy Tales (Challenge, 1998)
 * Zwart wit Bimhuis Live (BV Haast, 1999)
 * Umai (Challenge, 2000)
 * Brutto Gusto (Challenge, 2002)
 * Hidden History (Challenge, 2003)
 * Boompetit (Challenge, 2004)
 * Nocturnal Ghost Songs (Basta, 2005)
 * Summersault (Challenge, 2006)
 * The Compositions of Eric Dolphy with Alex Coke, Willem Breuker (BV Haast, 2006)
 * Gatecrashin'  (Challenge, 2007)
 * Hyper (Challenge, 2008)
 * Live at Yoshi's (Challenge, 2009)
 * Heavens Above! (Challenge, 2009)
 * Live at the Concertgebouw with Florian Weber (Challenge, 2011)
 * North Sea Jazz Legendary Concerts (Bob City, 2013)
 * Pessoa (Coast to Coast 2013)
 * Party Animals (V-Flow, 2020)

As sideman
With Michel Banabila
 * Urban Jazz Scapes (Javaanse Jongens, 2003)
 * Music for Viola and Electronics II (Tapu, 2015)
 * New Land (Tapu, 2017)

With Michiel Borstlap
 * The Sextet Live! (Challenge, 1995)
 * Body Acoustic (EmArcy 1999)
 * Liveline (EmArcy, 2000)

With Martin Fondse
 * Upperground (BV Haast, 1998)
 * Ere Ibeji (BV Haast, 2001)
 * Fragrant Moondrops (Basta, 2009)
 * Testimoni (Basta, 2012)

With others
 * Jan Akkerman, Minor Details (Digimode, 2011)
 * Jan Akkerman, North Sea Jazz Legendary Concerts (Bob City, 2013)
 * Kinan Azmeh, Jeroen Van Vliet, Eric Vloeimans' Levanter (V-Flow, 2018)
 * Bob Brookmeyer, Waltzing With Zoe (Challenge, 2001)
 * Calefax, On the Spot (Challenge, 2013)
 * Pierre Courbois, Live in Germany (A Records, 1996)
 * Pierre Courbois, Reouverture (A Records, 1996)
 * Freek de Jonge, Zonde (V2, 2013)
 * Rob de Nijs, Vanaf Vandaag (Capitol, 2004)
 * Chris Hinze, Senang (Keytone, 1996)
 * Brigitte Kaandorp, Thuis (Doek Op 2015)
 * Jacek Kochan, Another Blowfish (Not Two, 2002)
 * Michael Moore, Osiris (Ramboy, 2006)
 * Michael Moore, Rotterdam (Ramboy, 2011)
 * Enrico Pieranunzi, Alone Together (Challenge, 2001)
 * Ntjam Rosie, The One (Gentle Daze, 2015)
 * Florian Ross, Lilacs and Laughter (Naxos, 2000)
 * Sandhya Sanjana, Random Access Melody (LopLop, 2009)
 * Jarmo Savolainen, Grand Style (A Records, 2000)
 * Jarmo Savolainen, Times Like These (A Records, 2002)
 * Armin van Buuren, Embrace (Armada, 2015)
 * Jasper van 't Hof, Ballads of Timbuktu (Jaro Medien, 2002)
 * Jasper van 't Hof, NeverNeverLand (Jaro Medien, 2005)

title: Eric Vloeimans


text:
San Francisco Opera

The San Francisco Opera (SFO) is an American opera company founded in 1923 by Gaetano Merola (1881–1953) based in San Francisco, California.

Gaetano Merola (1923–1953)
Merola's road to prominence in the Bay Area began in 1906 when he first visited the city. In 1909, he returned as the conductor of the International Opera Company of Montreal, one of the many visiting troupes that frequented the bustling city. Continued visits over the next decade convinced him that an opera company in San Francisco was viable.

Merola moved back into the city in 1921 while living with Mrs. Oliver Stine's support Oliver Stine. He drafted plans for a new, locally-owned opera company that would not rely on visiting troupes, a common practice for some opera companies since the Gold Rush. By the next year, Merola organized a trial season at Stanford University. The first performance occurred in the Stanford Cardinal's football stadium on June 3, 1922, with operatic tenor Giovanni Martinelli performing in Pagliacci, Carmen, and Faust.

The five-day season's lacked any financial success; Merola instead successfully raised funds for a full season of opera in the following year. Following this, the first unaffiliated performance given by the San Francisco Opera was La bohème, with Queena Mario and Giovanni Martinelli, on September 26, 1923, in the city's Civic Auditorium.

In subsequent years, SFO would perform a wide array of Italian operas, rarely performing a given opera more than once. SFO seasons seldom lasted more than two months, and occasionally performed less than one month. The 1923-24 season included productions of Andrea Chénier and Mefistofele with Beniamino Gigli, Tosca with Giuseppe De Luca and Martinelli, and Verdi's Rigoletto with Queena Mario, De Luca, and Gigli.

During the nine years following the opening season, the San Francisco War Memorial Opera House was designed by Arthur Brown Jr. On October 15, 1932, the company inaugurated the new opera house with a performance of Tosca on with Claudia Muzio in the titular role. Characteristic of Merola's years as general director was the fact (as noted by Chatfield-Taylor) that "the great singers of the world came regularly to San Francisco, often performing several roles in deference to the short season and long travel time across the country."

Edwin MacArthur led the San Francisco Opera Orchestra in several 78-rpm recordings for RCA Victor in the late 1930s, including performances by soprano Kirsten Flagstad. Some of these were later reissued by RCA on LP and CD.

Short versions of all the works in the season were broadcast on about 30 California, Oregon, Washington, Idaho and British Columbia radio stations, starting about 1941.

Merola's tenure lasted 30 years. Characteristics of Merola's SFO included creating opportunities for young American singers regardless of the fact that the opera lacked a formal training program at that time, and also regular tours by the SFO to Los Angeles between 1937 and 1965, which expanded the season into November. However, until well after Merola's death on August 30, 1953, while conducting an open-air concert at Stern Grove, the main San Francisco season rarely extended beyond late October.

Kurt Herbert Adler (1953–1981)
Kurt Herbert Adler (1905–1988) came to the United States in 1938 after early experience and training in many aspects of music and theatre in Austria, Germany, and Italy. For five years, he worked to build the chorus of the Chicago Opera Company. Merola heard of him and, over the telephone, invited him to San Francisco opera in 1943 as chorus director. Adler was often regarded as a difficult, sometimes tyrannical person to work for. However, as Chatfield-Taylor notes, "singers, conductors, directors, and designers came back season after season. They came back because Adler made the SFO an internationally respected company that ran at a high level of professionalism and offered them interesting things to do in a warm and supportive atmosphere." Among those who were offered new and exciting challenges were Geraint Evans, the Welsh baritone, Leontyne Price, and Luciano Pavarotti.

He took on more and more administrative details as Merola's health and energy diminished, but Adler was not the board's natural choice to replace Merola at the time of his death in 1953. After three months of acting as artistic director, and with the assistance of its president, Robert Watt Miller, Adler was confirmed as general director.

Adler's aims
Adler's had several aims in taking over the company. One was to expand the season which in Merola's time ran. It changed from the Friday after Labor Day until early November (when the Metropolitan Opera's season began). This was to capitalize on the availability of singers by presenting up to fourteen operas with two or three performances each. Eventually, as seen in the 1961 SFO season, eleven operas were given five or six performances each on average while the season ran to late November.

Another aim was to present new talent, and, for this, he was tireless in seeking out up-and-coming new singers, whether American or European, by attending performances in both major and minor opera houses. He heard Leontyne Price on the radio and offered her a role in Dialogues of the Carmelites in 1957, thus providing her with her first performance on a major operatic stage. A short time later in the same season, she was to step into the role of Aida at short notice to replace Antonietta Stella, a role which gave her long-lived international acclaim.

Adler developed a strong professional relationship with Birgit Nilsson. Nilsson made her US-debut in San Francisco in 1956. She returned almost every season for 26 years and made her last appearance in the house in 1982.

A characteristic of the Adler years was the interest in developing stronger connections to opera stage directors in an attempt to strengthen the dramatic and theatrical elements of the works. In this, he was greatly supported by his long relationship with Jean-Pierre Ponnelle, the often-controversial stage director and designer who began his association with SFO in 1957.

Merola Opera Program
Several innovations undertaken by Adler included the Merola Opera Program (named after the first general director). It began during the 1954/55 Season and was given its current name in 1957. The program now annually offers approximately 23 gifted singers, four apprentice coaches, and one apprentice stage director the rare opportunity of studying, coaching, and participating in master classes with established professionals for eleven weeks during the summer. Many went on to international careers, among them Carol Vaness and Thomas Hampson.

Opera in the Park
Another innovation was "Opera in the Park" which, since 1971, has been an annual free concert in Golden Gate Park on the Sunday following opening night of the Fall Season. It traditionally features artists from the opening weekend in full concert with the San Francisco Opera Orchestra. The event is open to the public and draws some 30,000 listeners. The concert is presented in conjunction with the non-profit San Francisco Parks Trust and the San Francisco Chronicle Charities.

Success of the company
By the 1970s, the company was highly successful and offered audiences the "cream of the crop" of internationally known singers, but with Adler often bringing in unknowns to make their American debuts or the surprise of well-known singers replacing ailing ones, there were some exciting nights at the opera. These included Plácido Domingo flying with no notice from New York City to San Francisco – albeit three hours after curtain time – to replace the ailing Carlo Cossutta on the opening night of Otello and the last-minute substitution by Leontyne Price for Margaret Price in the role of Aida.

From 1971 to 1979, San Francisco station KKHI broadcast the regular Friday night performances of the opera on AM and FM (in multiplex stereo with quadraphonic encoding). The broadcasts were hosted by several well-known announcers, including Scott Beach and Fred Cherry.

In the summer of 1972, the San Francisco Opera began its 50th anniversary celebrations with a special free concert in Sigmund Stern Grove. Adler conducted most of the program, which featured performances by many of the surviving singers who had appeared with the company during its history. The legendary tenor Lauritz Melchior conducted the orchestra, rather than sing, in a performance of the famous Radetsky March by Johann Strauss I; it was possibly his last public appearance. One of the highlights of the afternoon program was a moving performance of the love duet from Madama Butterfly with soprano Licia Albanese and tenor Frederick Jagel.

Adler retired on December 15, 1981.

Terence McEwen (1982–1988)
Following Adler's retirement announcement in June 1979, Terence A. McEwen (1929–1998) was Adler's hand-picked successor. Growing up in the Montreal area of Canada, McEwen learned to love opera at an early age, listened to the Met broadcasts, and at age 14, made a trip to New York one winter break to hear several of his favorite operas, which included Bidu Sayão and Jussi Björling in Rigoletto. As a singer, Sayão was forever to remain his passion, one which was accentuated by seeing her in Manon performances in Montreal.

His passion for opera in general led him to visit the Royal Opera House in London and a lowly paid job with Decca Records in that city. Moving up the ranks in the 1950s, he landed in New York in 1959 and for the next 20 years made London Records, Decca's classical arm, the most significant classical label in the United States.

After being approached by Adler regarding the San Francisco opera job, he moved to the city in 1980 and involved himself totally in learning the running of an opera company. In January 1982 McEwen was running the SFO.

Given his expertise and background in understanding opera and the wonders of the human voice, it is not surprising that his approach in his early years was away from the theatrical side and more focused on singers. With his Ring Cycle which began in the Summer 1983 and Fall 1984 seasons — and which was presented in its entirety in June 1985 – McEwen demonstrated where his priorities lay: they were focused on hiring the best singers in the world.

As a reaction to the economic climate of the times, in 1982 McEwen, created the "San Francisco Opera Center" to oversee and combine the operation and administration of the numerous affiliate educational and training programs. Providing a coordinated sequence of performance and study opportunities for young artists, the San Francisco Opera Center included the "Merola Opera Program", "Adler Fellowship Program", "Showcase Series", "Brown Bag Opera", "Opera Center Singers", "Schwabacher Recitals", and various Education Programs. By introducing his young singers to the great voices of the past, inviting them to rehearsals, and giving tickets to current productions McEwen hoped to create rounded performers who could appear in the regular Fall season.

Among his successes in this regard was the mezzo-soprano Dolora Zajick from Nevada. By "hand holding" her through the various stages of training, he prepared her for the role of Azucena in Il trovatore for the summer 1986 season to great acclaim.

During the 1983 fall season, the student/family matinee performances of La traviata were presented with supertitles. These are English translations of the libretto projected over the proscenium simultaneously with the action on stage. The overwhelmingly favorable response prompted the company to introduce the practice in increasing numbers of performances in subsequent seasons. Supertitles are now used for all San Francisco Opera productions and SFO also rents its supertitles internationally to other opera companies.

In 1986, Sir John Pritchard was appointed music director and served until 1989.

On 8 February 1988, McEwen announced his resignation. The following day his mentor, Kurt Herbert Adler, died.

Lotfi Mansouri (1988–2001)
Lotfi Mansouri (born 1929) was already a known quantity when Terry McEwen announced his retirement. Then head of the Canadian Opera Company in Toronto, Mansouri had received an education in medicine in Los Angeles, but gave it all up upon becoming fixated on opera, first as a young tenor with UCLA's Opera Workshop, and then with opera in general.

As early as 1962, with Mansouri having found work as director in Los Angeles followed by his becoming resident stage director at the Zürich Opera, Adler came to see him at work, and he was offered six operas to direct in the 1963 season. By the time he became general director, he had directed 60 operas for SFO and many others elsewhere.

By 1975 he was director of the Canadian Opera Company where, in 1983, he had introduced the revolutionary supertitles. Mansouri's feelings on the effects of titling was that the audience would be more engaged in the performance. This was a momentous change in the world of opera.

Mansouri introduced many new operas to the SFO repertory. These included more Russian operas with the highlight being Valery Gergiev's conducting of Prokofiev's War and Peace and a firm link established with the Kirov Opera. Also, there was Rossini's Guillaume Tell and Verdi's I vespri siciliani which followed.

One of Mansouri's triumphs was the overseeing of the reconstruction and renovation of the opera house following the October 1989 earthquake. After closing at the end of the 1995 Fall season for "a 21-month, US$88.5 million renovation, San Francisco's War Memorial Opera House reopened on 5 September 1997 with a gala concert celebrating this occasion, as well the 75th anniversary of the San Francisco Opera. Fittingly, the concert featured operatic greats of the past, present and future. The project included repairs of damage caused by the 1989 Loma Prieta earthquake, improvements for the audience and performers, seismic strengthening and a general cleanup that left the 65-year-old Opera House gleaming."

Donald Runnicles was named music director and principal conductor of SFO in 1990, and assumed the posts in 1992.

In November 1992, Mansouri introduced "Pacific Visions", an ambitious program designed to maintain the vitality of the opera repertoire through new commissions and the presentation of unusual repertoire. It was launched with the commissioning of the following operas:

 * The Dangerous Liaisons, composed by Conrad Susa to a libretto by Philip Littell, had its world premiere during the 1994 fall season and was the subject of a nationwide TV broadcast. The cast featured Renée Fleming, Frederica von Stade, and Thomas Hampson.
 * Harvey Milk, composer Stewart Wallace to a libretto by Michael Korie. It was performed in 1996 as a joint commission and co-production of the SFO, Houston Grand Opera, and New York City Opera. The cast featured Raymond Very as Dan White, Robert Orth as Harvey Milk and Gidon Sachs as Mayor Moscone.
 * A Streetcar Named Desire, composed by André Previn to a libretto by Philip Littell, after the play by Tennessee Williams. The work had its premiere during the 1998–99 fall season. The cast included Renée Fleming as Blanche DuBois and Elizabeth Futral as Stella, baritone Rod Gilfry as Stanley Kowalski and tenor Anthony Dean Griffey as Mitch.
 * Dead Man Walking, composed by Jake Heggie from a libretto by Terrence McNally after the book by Sister Helen Prejean, received its premiere in October 2000. The cast included Susan Graham as Sister Helen Prejean, John Packard as Joe; and Frederica von Stade as Mrs. Patrick de Rocher.
 * The Death of Klinghoffer(a co-commission between San Francisco Opera, Brussels' La Monnaie, the Opera de Lyon, the Los Angeles Festival, the Glyndebourne Festival and the Booklyn Academy of Music), composed by John Adams in 1992. The cast included Janice Felty in 3 roles, James Maddalena as The Captain, and Thomas Hammons as the First Officer.

Summing up his years at the SFO, the San Francisco Chronicle noted: "He's never been interested in the succès d'estime, the daring intellectual or theatrical coup that dazzles culture mavens but leaves the general public alienated or bewildered. For Mansouri, a success that doesn't put fannies in the seats is no success at all."

Towards the end of the 2001 season, Mansouri announced his retirement after fourteen seasons with SFO and 50 years in opera.

Pamela Rosenberg (2001–2005)
's first connection with the San Francisco Opera was as a standee while attending the University of California, Berkeley. She returned to SFO with a background of operatic productions in Germany and, specifically, as head of the Stuttgart Opera.

In January 2001, Rosenberg announced her first artistic initiative for San Francisco Opera, "Animating Opera", a multi-year plan of interwoven themes and series. These included "Seminal Works of Modern Times", "The Faust Project", "Composer Portrait: Janáček/Berlioz", "Women Outside of Society: Laws Unto Themselves", "Metamorphosis: From Fairy Tales to Nightmares", and "Outsiders or Pioneers?: The Nature of the Human Condition".

Incorporated within the production programming of "Animating Opera" was the America staged premiere of Messiaen's Saint-François d'Assise, Virgil Thomson's The Mother of Us All, as well as a commission for a new work by John Adams and Peter Sellars entitled Doctor Atomic, which premiered on 1 October 2005. Other operas new to the SFO's repertoire during her directorship include Busoni's Doktor Faust, Ligeti's Le Grand Macabre and Janáček's The Cunning Little Vixen.

After much controversy surrounding her management of the SFO, which included deficits created after the "dot-com" collapse in 2000 and the effects of September 11 on arts attendance, she announced in 2004 that she would not renew her contract with the company when it ended in late 2005.

As noted by Steven Winn in the San Francisco Chronicle in December 2005, "Productions were scuttled or postponed in the face of a $7.7 million deficit. Ambitious programming initiatives and plans for a second, smaller performance venue went by the wayside. Company-wide cuts pared 14 percent from the company's $67 million budget in 2003."

He continued: "Embattled by financial woes and trying labor negotiations, Rosenberg was routinely blamed for problems that were largely beyond her control. Her taste for new and unusual operas and a European-honed aesthetic that favored brash and even radical reinterpretations of the classics, the thinking went, drove away audiences and donors and ran up costs in the company's hour of greatest need."

Rosenberg returned to Germany to work with Sir Simon Rattle and the Berlin Philharmonic as its Intendantin. From 2004 to 2007, Keith Cerny served as the chief financial officer of the San Francisco Opera.

David Gockley (2006–2016)
After 33 years of directing the Houston Grand Opera, David Gockley became the SFO's General Director on 1 January 2006. As part of an announcement of the 2006/2007 season and the future of the company on 11 January, Gockley noted that "this season we debut a new visual identity and logo in keeping with a new artistic philosophy. I believe that it speaks of glamour, sophistication, tradition and innovation all things that infuse our plans for the future of San Francisco Opera."

In May 2011, it was announced that Gockley's contract was to be extended through SFO's 2015–16 season. In October 2014 it was announced that Gockley would be stepping down from his post at the end of the 2015/16 season. His replacement was announced to be Matthew Shilvock in September 2015.

Innovations announced
As part of his future plans, Gockley stated "I want nothing less than to have the greatest stars of the opera world perform here regularly. You can expect in coming seasons to hear Renée Fleming, Anna Netrebko, Thomas Hampson, Dmitri Hvorostovsky, Marcello Giordani, Ramón Vargas, Marcelo Álvarez, Juan Diego Flórez, Ben Heppner, Natalie Dessay, and Angela Gheorghiu, among many others. We will have a world premiere for you in 2007, and the Wagner lovers among you will be happy to hear that we expect to commence a Ring Cycle in 2008."

Ring Cycle

San Francisco Opera and Washington National Opera began a co-production of a new Ring Cycle in 2006 directed by Francesca Zambello. The production used imagery from various eras of American history and had a feminist and environmentalist viewpoint. SFO presented Das Rheingold in June 2008, Die Walküre in June 2010, and three complete Ring cycles in June 2011. The complete cycles in June 2011 were conducted by Donald Runnicles and featured cycle role debuts of Mark Delavan (Wotan) and Nina Stemme (Brünnhilde) as well as Jay Hunter Morris (making his role debut in the title role of Siegfried) and Ian Storey (making his role debut as Siegfried in Götterdämmerung.

Technological innovations

In May 2006 Gockley oversaw SFO's first simulcast, a live broadcast of a mainstage performance of Madama Butterfly to San Francisco's Civic Center Plaza for a crowd of 8,000. Subsequent simulcasts have been presented at Stanford University's Frost Amphitheater, four theaters in Northern California, and San Francisco's AT&T Park. SFO's has simulcast nine operas to AT&T Park since 2007 that have collectively drawn more than 165,000 opera fans.

The technology for the simulcasts and other innovations like OperaVision—a series of screens located throughout the War Memorial Opera House that project close-up shots of the action on stage—is made possible through SFO's Koret-Taube Media Suite. Completed in 2007, The Koret-Taube Media Suite is the first permanent high-definition broadcast-standard video production facility installed in any American opera house according to the company's website.

In 2007, San Francisco Opera returned to regular broadcasts of its productions on national and international radio., and in December the Opera announced the presentation of four operas in movie theaters across the United States. Following the initial presentation of the four operas in movie theaters in 2008, San Francisco Opera used these four titles to create its Grand Opera Cinema Series, making these titles available to be presented by performing arts centers, theaters, and universities. Since 2008 the company has added eight additional operas to the Grand Opera Cinema Series, and they have been presented by KQED-TV with hosts Rita Moreno and Joan Chen.

Music directors and conductors under Gockley
In September 2006, it was announced and reported that by mutual agreement with Gockley, Donald Runnicles would conclude his tenure as music director in 2009. However, he has maintained an association with SFO and conducted the 2010/11 production of Der Ring des Nibelungen as well as 2015's Les Troyens.

On 9 January 2007, SFO announced its third music director would be the Italian conductor Nicola Luisotti, beginning with the 2009/10 season, for an initial contract of 5 years. Luisotti made his SFO debut in 2005 with La forza del destino, and returned in 2008 to conduct La bohème prior to assuming the role of music director. In SFO's September 2009 program magazine, David Gockley announced that bringing on Luisotti as the company's music director was a large part of his goal to "reinvigorate the core Italian repertory that is San Francisco Opera's birthright." Gockley also stated that Luisotti would conduct three to four productions each season, including one non-Italian opera; since 2009 these non-Italian operas have included Salome, Lohengrin, and Carmen.

In January 2009, Gockley announced the reappointment of Patrick Summers as principal guest conductor and named Giuseppe Finzi as the company's new assistant music director. Finzi was named as SFO's resident conductor in 2011.

Programming
The company secured composer John Adams' Nixon in China for their 2011/12 season.

San Francisco Opera has presented several world premieres under David Gockley's tenure. So far these include Philip Glass and Christopher Hampton's Appomattox in 2007; Stewart Wallace and Amy Tan's The Bonesetter's Daughter in 2008; and Christopher Theofanidis and Donna Di Novelli's Heart of a Soldier in 2011. In 2013, the company presented three world premieres: Nolan Gasser and Carey Harrison's The Secret Garden, based on the children's book by Frances Hodgson Burnett (and staged in conjunction with UC Berkeley's Cal Performances); Mark Adamo's The Gospel of Mary Magdalene; and Tobias Picker and J. D. McClatchy's Dolores Claiborne, based on the novel by Stephen King. In the summer of 2015 the world premiere of La Ciociara by Marco Tutino and Luca Rossi, based on novel of the same name by Alberto Moravia, took place.

San Francisco Opera's recent commissions include Dream of the Red Chamber by Bright Sheng and David Henry Hwang in the fall of 2016, based on the work of the same name by 18th-century Qing Dynasty writer Cao Xueqin.

Wilsey Center for Opera
In order to consolidate its various office and work spaces scattered throughout San Francisco, SFO took over the fourth floor of the War Memorial and Performing Arts Center in February 2016, following the completion of the building's $156-million seismic retrofit and renovation. To accomplish this, the company started a campaign to name various locations of the new space after donors in 2011. San Francisco philanthropist Dede Wilsey pledged a $5-million gift, and the entire facility was named the Diane B. Wilsey Center for Opera. Designed by San Francisco architectural firm Mark Cavagnero Associates, the center provides additional office space as well as costume storage; two multipurpose rooms for rehearsals, board meetings, and social events; and a 299-seat performance venue.

title: San Francisco Opera