Lactarius indigo

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Lactarius indigo
Lactarius indigo 48568 edit.jpg
The gills of L. indigo
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Russulales
Family: Russulaceae
Genus: Lactarius
Species:
L. indigo
Binomial name
Lactarius indigo
(Schwein.) Fr. (1838)
Synonyms

Agaricus indigoSchwein. (1822)
Lactarius canadensisWinder (1871) [1]
Lactifluus indigo(Schwein.) Kuntze (1891) [2]

Contents

Lactarius indigo
Information icon.svg
Gills icon.png Gills on hymenium
Depressed cap icon.svg Cap is depressed
Adnate gills icon2.svgDecurrent gills icon2.svg Hymenium is adnate or decurrent
Bare stipe icon.svg Stipe is bare
Transparent spore print icon.svg
Spore print is yellow
Mycorrhizal fungus.svgEcology is mycorrhizal
Mycomorphbox Edible.pngEdibility is edible

Lactarius indigo, commonly known as the indigo milk cap, indigo milky, the indigo (or blue) lactarius, or the blue milk mushroom, is a species of agaric fungus in the family Russulaceae. It is a widely distributed species, growing naturally in eastern North America, East Asia, and Central America; it has also been reported in southern France. L. indigo grows on the ground in both deciduous and coniferous forests, where it forms mycorrhizal associations with a broad range of trees. The fruit body color ranges from dark blue in fresh specimens to pale blue-gray in older ones. The milk, or latex, that oozes when the mushroom tissue is cut or broken — a feature common to all members of the genus Lactarius — is also indigo blue, but slowly turns green upon exposure to air. The cap has a diameter of 5–15 cm (2–6 in), and the stem is 2–8 cm (0.8–3 in) tall and 1–2.5 cm (0.4–1.0 in) thick. It is an edible mushroom, and is sold in rural markets in China, Guatemala, and Mexico. In Honduras, the mushroom is called a chora, and is generally eaten with egg; generally as a side dish for a bigger meal.

Taxonomy and nomenclature

Originally described in 1822 as Agaricus indigo by American mycologist Lewis David de Schweinitz, [3] the species was later transferred to the genus Lactarius in 1838 by the Swede Elias Magnus Fries. [4] German botanist Otto Kuntze called it Lactifluus indigo in his 1891 treatise Revisio Generum Plantarum , [2] but the suggested name change was not adopted by others. Hesler and Smith, in their 1960 study of North American species of Lactarius, defined L. indigo as the type species of subsection Caerulei, a group characterized by blue latex and a sticky, blue cap. [5] In 1979, they revised their opinions on the organization of subdivisions in the genus Lactarius, and instead placed L. indigo in subgenus Lactarius based on the color of latex and the subsequent color changes observed after exposure to air. [6] As they explained:

The gradual development of blue to violet pigmentation as one progresses from species to species is an interesting phenomenon deserving further study. The climax is reached in L. indigo which is blue throughout. L. chelidonium and its variety chelidonioides, L. paradoxus, and L. hemicyaneus may be considered as mileposts along the road to L. indigo. [7]

The specific epithet indigo is derived from the Latin word meaning "indigo blue". [8] Its names in the English vernacular include the "indigo milk cap", [9] the "indigo Lactarius", [8] the "blue milk mushroom", [10] and the "blue Lactarius". [11] In central Mexico, it is known as añil, azul, hongo azul, zuin, and zuine; it is also called quexque (meaning "blue") in Veracruz and Puebla. [12]

Description

Like many other mushrooms, L. indigo develops from a nodule that forms within the underground mycelium, a mass of threadlike fungal cells called hyphae that make up the bulk of the organism. Under appropriate environmental conditions of temperature, humidity, and nutrient availability, the visible reproductive structures (fruit bodies) are formed. The cap of the fruit body, measuring between 5–15 cm (2.0–5.9 in) in diameter, is initially convex and later develops a central depression; in age it becomes even more deeply depressed, becoming somewhat funnel-shaped as the edge of the cap lifts upward. [13] The margin of the cap is rolled inwards when young, but unrolls and elevates as it matures. The cap surface is indigo blue when fresh, but fades to a paler grayish- or silvery-blue, sometimes with greenish splotches. It is often zonate: marked with concentric lines that form alternating pale and darker zones, and the cap may have dark blue spots, especially towards the edge. Young caps are sticky to the touch. [14]

Lactarius indigo 54015.jpg
Overhead view of caps found in Guadalajara, Mexico
Lactarius indigo 48609.jpg
The cap margin is rolled inwards in young specimens.

The flesh is pallid to bluish in color, slowly turning greenish after being exposed to air; its taste is mild to slightly acrid. The flesh of the entire mushroom is brittle, and the stem, if bent sufficiently, will snap open cleanly. [15] The latex exuded from injured tissue is indigo blue, and stains the wounded tissue greenish; like the flesh, the latex has a mild taste. [8] Lactarius indigo is noted for not producing as much latex as other Lactarius species, [16] and older specimens in particular may be too dried out to produce any latex. [17]

The gills of the mushroom range from adnate (squarely attached to the stem) to slightly decurrent (running down the length of the stem), and crowded close together. Their color is an indigo blue, becoming paler with age or staining green with damage. The stem is 2–6 cm (0.8–2.4 in) tall by 1–2.5 cm (0.4–1.0 in) thick, and the same diameter throughout or sometimes narrowed at base. Its color is indigo blue to silvery- or grayish blue. The interior of the stem is solid and firm initially, but develops a hollow with age. [9] Like the cap, it is initially sticky or slimy to the touch when young, but soon dries out. [18] Its attachment to the cap is usually in a central position, although it may also be off-center. [19] Fruit bodies of L. indigo have no distinguishable odor. [20]

L. indigo var. diminutivus (the "smaller indigo milk cap") is a smaller variant of the mushroom, with a cap diameter between 3–7 cm (1.2–2.8 in), and a stem 1.5–4.0 cm (0.6–1.6 in) long and 0.3–1.0 cm (0.1–0.4 in) thick. [21] It is often seen in Virginia. [20] Hesler and Smith, who first described the variant based on specimens found in Brazoria County, Texas, described its typical habitat as "along [the] sides of a muddy ditch under grasses and weeds, [with] loblolly pine nearby". [22]

Microscopic features

When viewed in mass, as in a spore print, the spores appear cream to yellow colored. [8] [9] Viewed with a light microscope, the spores are translucent (hyaline), elliptical to nearly spherical in shape, with amyloid warts, and have dimensions of 7–9 by 5.5–7.5  μm. [8] Scanning electron microscopy reveals reticulations on the spore surface. [12] The hymenium is the spore-producing tissue layer of the fruit body, and consists of hyphae that extend into the gills and terminate as end cells. Various cell types can be observed in the hymenium, and the cells have microscopic characteristics that may be used to help identify or distinguish species in cases where the macroscopic characters may be ambiguous. The spore-bearing cells, the basidia, are four-spored and measure 37–45 μm long by 8–10 μm wide at the thickest point. [23] Cystidia are terminal cells of hyphae in the hymenium which do not produce spores, and function in aiding spore dispersal, and maintaining favorable humidity around developing spores. [24] The pleurocystidia are cystidia that are found on the face of a gill; they are 40–56 by 6.4–8 μm, roughly spindle-shaped, and have a constricted apex. The cheilocystidia—located on the edge of a gill—are abundant, and are 40.0–45.6 by 5.6–7.2 μm. [12]

The "milk", or latex, is indigo blue. Lactarius indigo 7283.jpg
The "milk", or latex, is indigo blue.

Similar species

The characteristic blue color of the fruiting body and the latex make this species easily recognizable. Other Lactarius species with some blue color include the "silver-blue milky" ( L. paradoxus ), found in eastern North America, [20] which has a grayish-blue cap when young, but it has reddish-brown to purple-brown latex and gills. L. chelidonium has a yellowish to dingy yellow-brown to bluish-gray cap and yellowish to brown latex. L. quieticolor has blue-colored flesh in the cap and orange to red-orange flesh in the base of the stem. [9] Although the blue discoloration of L. indigo is thought to be rare in the genus Lactarius, in 2007 five new species were reported from Peninsular Malaysia with bluing latex or flesh, including L. cyanescens , L. lazulinus , L. mirabilis , and two species still unnamed. [25]

Edibility

Although L. indigo is a well-known edible species, opinions vary on its desirability. For example, American mycologist David Arora considers it a "superior edible", [9] while a field guide on Kansas fungi rates it as "mediocre in quality". [26] It may have a slightly bitter, [27] or peppery taste, [28] and has a coarse, grainy texture. [8] [26] The firm flesh is best prepared by cutting the mushroom in thin slices. The blue color disappears with cooking, and the mushroom becomes grayish. Because of the granular texture of the flesh, it does not lend itself well to drying. Specimens producing copious quantities of milk may be used to add color to marinades. [29]

In Mexico, individuals harvest the wild mushrooms for sale at farmers' markets, typically from June to November; [12] they are considered a "second class" species for consumption. [30] L. indigo is also sold in Guatemalan markets from May to October. [31] It is one of 13 Lactarius species sold at rural markets in Yunnan in southwestern China. [32]

Chemical composition

(7-Isopropenyl-4-methylazulen-1-yl)methyl stearate, the blue lipophilic azulene pigment from L. indigo. The part of the molecule responsible for its color (chromophore) is highlighted in blue. Lactarius azulene.png
(7-Isopropenyl-4-methylazulen-1-yl)methyl stearate, the blue lipophilic azulene pigment from L. indigo. The part of the molecule responsible for its color (chromophore) is highlighted in blue.

A chemical analysis of Mexican specimens has shown L. indigo to contain moisture at 951 mg/g of mushroom, fat at 4.3 mg/g, protein at 13.4 mg/g, and dietary fiber at 18.7 mg/g, much higher in comparison to the common button mushroom, which contains 6.6 mg/g. Compared to three other wild edible mushroom species also tested in the study ( Amanita rubescens , Boletus frostii , and Ramaria flava ), L. indigo contained the highest saturated fatty acids content, including stearic acid with 32.1 mg/g—slightly over half of the total free fatty acid content. [33]

The blue color of L. indigo is due to (7-isopropenyl-4-methylazulen-1-yl)methyl stearate, an organic derivative of azulene, which is biosynthesised from a sesquiterpene very similar to matricin, the precursor for chamazulene. It is unique to this species, but similar to a compound found in L. deliciosus . [34]

Distribution, habitat, and ecology

Lactarius indigo is distributed throughout southern and eastern North America but is most common along the Gulf Coast, Mexico, and Guatemala. Its frequency of appearance in the Appalachian Mountains of the United States has been described as "occasional to locally common". [8] Mycologist David Arora notes that in the United States, the species is found with ponderosa pine in Arizona, but is absent in California's ponderosa pine forests. [35] It has also been collected from China, [32] India, [36] [37] Guatemala, [38] Costa Rica (in forests dominated by oak), [39] and as its southernmost distribution in the Humboldt oak cloud forests of Colombia. [40] In Europe, it has so far only been found in southern France. [41] A study on the seasonal appearance of fruiting bodies in the subtropical forests of Xalapa, Mexico, confirmed that maximal production coincided with the rainy season between June and September. [42]

A collection of L. indigo found in Jalisco, Mexico Lactarius indigo 19700.jpg
A collection of L. indigo found in Jalisco, Mexico

L. indigo is a mycorrhizal fungus, and as such, establishes a mutualistic relationship with the roots of certain trees ("hosts"), in which the fungi exchange minerals and amino acids extracted from the soil for fixed carbon from the host. The subterranean hyphae of the fungus grow a sheath of tissue around the rootlets of a broad range of tree species, forming so-called ectomycorrhizae—an intimate association that is especially beneficial to the host, as the fungus produces enzymes that mineralize organic compounds and facilitate the transfer of nutrients to the tree. [43]

Reflecting their close relationships with trees, the fruit bodies of L. indigo are typically found growing on the ground, scattered or in groups, in both deciduous and coniferous forests. [44] They are also commonly found in floodplain areas that have been recently submerged. [17] In Mexico, associations have been noted with Mexican alder, American Hornbeam, American Hophornbeam, and Liquidambar macrophylla, [12] while in Guatemala the mushroom associates with smooth-bark Mexican pine and other pine and oak species. [38] In Costa Rica, the species forms associations with several native oaks of the genus Quercus . [45] Under controlled laboratory conditions, L. indigo was shown to be able to form ectomycorrhizal associations with the neotropical pine species Mexican white pine, Hartweg's pine, Mexican yellow pine, smooth-bark Mexican pine, [31] and the Eurasian pines Aleppo pine, European black pine, maritime pine, and Scots pine. [46]

See also

Related Research Articles

<i>Lactarius</i> Genus of fungi

Lactarius is a genus of mushroom-producing, ectomycorrhizal fungi, containing several edible species. The species of the genus, commonly known as milk-caps, are characterized by the milky fluid ("latex") they exude when cut or damaged. Like the closely related genus Russula, their flesh has a distinctive brittle consistency. It is a large genus with over 500 known species, mainly distributed in the Northern hemisphere. Recently, the genus Lactifluus has been separated from Lactarius based on molecular phylogenetic evidence.

<i>Lactarius deterrimus</i> Species of fungus

Lactarius deterrimus, also known as false saffron milkcap or orange milkcap, is a species of fungus in the family Russulaceae. The fungus produces medium-sized fruit bodies (mushrooms) with orangish caps up to 12 centimetres wide that develop green spots in old age or if injured. Its orange-coloured latex stains maroon within 30 minutes. Lactarius deterrimus is a mycorrhizal fungus that associates with Norway spruce and bearberry. The species is distributed in Europe, but has also found in parts of Asia. A visually similar species in the United States and Mexico is not closely related to the European species. Fruit bodies appear between late June and November, usually in spruce forests. Although the fungus is edible—like all Lactarius mushrooms from the section Deliciosi—its taste is often bitter, and it is not highly valued. The fruit bodies are used as source of food for the larvae of several insect species. Lactarius deterrimus can be distinguished from similar Lactarius species by difference in the mycorrhizal host or latex colour.

<i>Lactarius deliciosus</i> Species of fungus

Lactarius deliciosus, commonly known as the delicious milk cap, saffron milk cap and red pine mushroom, is one of the best known members of the large milk-cap genus Lactarius in the order Russulales. It is native to Europe, but has been accidentally introduced to other countries along with pine trees, with which the fungus is symbiotic.

<i>Lactarius torminosus</i> Fungus in the family Russulaceae from North Africa, northern Asia, Europe, and North America

Lactarius torminosus, commonly known as the woolly milkcap or the bearded milkcap, is a large agaric fungus. A common and widely distributed species, it is found in North Africa, northern Asia, Europe, and North America. It was first described scientifically by Jacob Christian Schäffer in 1774 as an Agaricus, and later transferred to the genus Lactarius in 1821 by Samuel Frederick Gray. A variety, L. torminosus var. nordmanensis, is known from the United States, Canada, and Switzerland. L. torminosus officially became the type species of Lactarius in 2011 after molecular studies prompted the taxonomic reshuffling of species between several Russulaceae genera.

<span class="mw-page-title-main">Candy cap</span> Species of fungus

Candy cap or curry milkcap is the English-language common name for several closely related edible species of Lactarius; L. camphoratus, L. fragilis, and L. rubidus. These mushrooms are valued for their highly aromatic qualities and are used culinarily as a flavoring rather than as a constituent of a full meal.

<i>Lactarius rubrilacteus</i> Species of fungus

Lactarius rubrilacteus is a species of mushroom of the genus Lactarius. It is also known as the bleeding milkcap, as is at least one other member of the genus, Lactarius sanguifluus.

<i>Lactifluus volemus</i> Species of edible fungus in the family Russulaceae widely distributed in the northern hemisphere

Lactifluus volemus, formerly known as Lactarius volemus, and commonly known as the weeping milk cap or bradley, is a species of fungus in the family Russulaceae. It is widely distributed in the northern hemisphere, in temperate regions of Europe, North America and Asia as well as some subtropical and tropical regions of Central America and Asia. A mycorrhizal fungus, its fruit bodies grow on the ground at the base of various species of trees from summer to autumn, either individually or in groups. It is valued as an edible mushroom, and is sold in markets in Asia. Several other Lactifluus mushrooms resemble L. volemus, such as the closely related edible species L. corrugis, but these can be distinguished by differences in distribution, visible morphology, and microscopic characteristics. L. volemus produces a white spore print and has roughly spherical spores about 7–8 micrometres in diameter.

<i>Lactarius subflammeus</i> Species of fungus

Lactarius subflammeus, commonly known as the orange milk cap, is a species of fungus in the family Russulaceae. It is found in western North America in the late summer and fall and is especially common in the Pacific Northwest, where it grows on the ground near conifers like pine and spruce. The brightly colored fruit bodies, which are slimy or sticky, have scarlet caps when young that soon fade to brilliant orange. The stem—typically longer than the width of the cap—is also bright orange but the gills are whitish. The mushroom secretes a whitish latex when it is cut or injured.

<i>Lactarius aspideoides</i> Species of fungus

Lactarius aspideoides, commonly known as the bright yellow milkcap, is a species of fungus in the family Russulaceae. Distinctive characteristics of the fruit body include the sticky, pale yellow cap, and the abundant latex that stains the mushroom tissue lilac. It is found in North America.

<i>Lactarius alnicola</i> Species of fungus

Lactarius alnicola, commonly known as the golden milkcap, is a species of fungus in the family Russulaceae. The fruit bodies produced by the fungus are characterized by a sticky, vanilla-colored cap up to 20 cm (7.9 in) wide with a mixture of yellow tones arranged in faint concentric bands. The stem is up to 5 cm (2.0 in) long and has yellow-brown spots. When it is cut or injured, the mushroom oozes a white latex, which has an intensely peppery taste. The acrid taste of the fruit bodies renders them unpalatable. The fungus is found in the western United States and Mexico, where it grows in mycorrhizal associations with various coniferous trees species, such as spruce, pine and fir, and deciduous species such as oak and alder. It has also been collected in India. Two varieties have been named: var. pitkinensis, known from Colorado, and var. pungens, from Michigan.

<i>Lactarius pubescens</i> Species of fungus

Lactarius pubescens, commonly known as the downy milk cap, is a species of fungus in the family Russulaceae. It is a medium to large agaric with a creamy-buff, hairy cap, whitish gills and short stout stem. The fungus has a cosmopolitan distribution, and grows solitarily or in scattered groups on sandy soil under or near birch.

<i>Lactarius repraesentaneus</i> Species of fungus

Lactarius repraesentaneus, commonly known as the northern bearded milkcap, the northern milkcap, or the purple-staining bearded milkcap, is a species of fungus in the family Russulaceae. It has a northerly distribution, and is found in temperate regions of North America and Europe, associated with spruce trees. Distinguishing features of its fruit body include the large orange-yellow cap up to 18 cm (7.1 in) wide, cream to pale yellow gills, and a yellow coarsely-pitted stem that is up to 12 cm (4.7 in) long and 3 cm (1.2 in) thick. Cut fruit bodies ooze a white latex that will stain mushroom tissue lilac to purple. Several chemicals have been isolated and identified from the fruit bodies that can modify the growth of plants, and the mushroom also has antibiotic activity against Staphylococcus aureus. L. repraesentaneus is poisonous, and consumption causes stomach aches.

<i>Lactarius affinis</i> Species of mushroom

Lactarius affinis, commonly known as the kindred milk cap, is a species of milk-cap mushroom in the family Russulaceae. It is found northeastern North America, where it fruits in the summer and fall, and is common in the Great Lakes region. Its fruit bodies have medium to large, slimy dull yellow or brownish caps. Although not considered poisonous, it is unpalatable because of its highly acrid taste.

<i>Lactarius fumosus</i> Species of fungus

Lactarius fumosus, commonly known as the smoky milkcap, is a species of fungus in the family Russulaceae.

<i>Lactarius fallax</i> Species of fungus

Lactarius fallax, commonly known as the velvety milk cap, is a species of fungus in the family Russulaceae. Found in both spruce and mixed conifer forests, it is a fairly common species in the Pacific Northwest region of North America, with a northerly range extending to Alaska. Its fruit bodies are medium-sized, with velvety, brown to blackish caps up to 3–9 cm (1.2–3.5 in) in diameter bearing a distinct pointed umbo. The caps are supported by velvety stems up to 6 cm (2.4 in) long and 1.5 cm (0.6 in) thick. The mushroom oozes a whitish latex when it is cut, and injured tissue eventually turns a dull reddish color. The eastern North American and European species Lactarius lignyotus is closely similar in appearance, but can be distinguished by its differing range.

<span class="mw-page-title-main">Lactifluus deceptivus</span> Species of fungus

Lactifluus deceptivus, commonly known as the deceiving milkcap, is a common species of fungus in the family Russulaceae. It is found throughout eastern North America on the ground in coniferous forests near hemlock or deciduous forests near oak, and in oak-dominated forests of Costa Rica. It produces large mushrooms with funnel-shaped caps reaching up to 25 cm (9.8 in) in diameter, on top of hard white stems that may reach 4–10 cm (1.6–3.9 in) long and up to 3 cm (1.2 in) thick. The gills are closely spaced together and yellowish-cream in color. When young, the cap is white in all parts, but the depressed center becomes dull brownish in age and breaks up into scales. The edge of the cap has a roll of cottony tissue that collapses as the cap expands. The surface of the stem—especially near the base—has a velvety texture. The mushroom "bleeds" a milky white acrid latex when it is cut or injured. Similar milk-cap species with which L. deceptivus might be confused include Lactifluus piperatus, L. pseudodeceptivus, L. caeruleitinctus, L. subvellereus, Lactarius arcuatus and Lactarius parvulus.

<i>Lactarius argillaceifolius</i> Species of fungus

Lactarius argillaceifolius is a species of fungus in the family Russulaceae. The mushrooms produced by the fungus have convex to flattened drab lilac-colored caps that are up to 18 cm (7.1 in) wide. The cream-colored gills are closely spaced together and extend slightly down the length of the stem, which is up to 9 cm (3.5 in) long by 3.5 cm (1.4 in) thick. The mushroom produces an off-white latex when injured that stains the mushroom tissue brownish.

<i>Lactarius rufulus</i> Species of fungus

Lactarius rufulus, commonly known as the rufous candy cap, is a species of fungus in the family Russulaceae. The fruit bodies have fleshy brownish-red caps up to 10 cm (3.9 in) wide, and closely spaced pinkish-yellow gills. The stem is up to 12 cm (4.7 in) long and 3 cm (1.2 in) thick and colored similarly to the cap. The species, known only from California, Arizona, and Mexico, grows on the ground in leaf litter near oak trees. The fruit bodies resembles those of L. rufus, but L. rufulus tends to grow in clusters at a common base, rather than solitarily or in groups. A distinguishing microscopic characteristic is the near absence of large, spherical cells called sphaerocysts that are otherwise common in Lactarius species. Lactarius rufulus mushrooms are edible, and have an odor resembling maple syrup. They have been used to flavor confections and desserts.

<i>Lactarius vinaceorufescens</i> Species of fungus

Lactarius vinaceorufescens, commonly known as the yellow-staining milkcap or the yellow-latex milky, is a poisonous species of fungus in the family Russulaceae. It produces mushrooms with pinkish-cinnamon caps up to 12 cm (4.7 in) wide held by pinkish-white stems up to 7 cm (2.8 in) long. The closely spaced whitish to pinkish buff gills develop wine-red spots in age. When it is cut or injured, the mushroom oozes a white latex that rapidly turns bright sulfur-yellow. The species, common and widely distributed in North America, grows in the ground in association with conifer trees. There are several other Lactarius species that bear resemblance to L. vinaceorufescens, but most can be distinguished by differences in staining reactions, macroscopic characteristics, or habitat.

<i>Lactarius pseudomucidus</i> Species of fungus

Lactarius pseudomucidus, commonly known as the slimy milk cap, is a fungus native to the northwestern part of North America, often found in coastal and conifer forests. It has a charcoal brown cap, smooth and slimy, from 2–10 cm across, initially flat convex, becoming shallowly depressed. The gills are decurrent, white with a gray or yellow tinge, staining brownish. The stipe is 40–100 mm tall, hollow, brittle. Both the cap and stipe are mucilaginous. The flesh is gray and the latex is milky white, drying yellowish. There is only a slight odor, and the taste slowly becomes acrid. Spores are white in mass, ellipsoid, amyloid, about 8 μm long, with a reticulate decoration on the surface. The species is inedible. It resembles Lactarius argillaceifolius, which has a light orange-gray cap, and eastern North America's Lactarius mucidus. It's edibility is unknown, but the extremely viscid stalk and cap are a deterrent.

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