Morchella esculenta

Last updated

Morchella esculenta
Morchella esculenta - DE - TH - 2013-05-01 - 01.JPG
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Pezizomycetes
Order: Pezizales
Family: Morchellaceae
Genus: Morchella
Species:
M. esculenta
Binomial name
Morchella esculenta
Fr.
Synonyms

Helvella esculenta(L.) Sowerby
Phallus esculentusL.

Contents

Morchella esculenta
Information icon.svg
Smooth icon.pngSmooth hymenium
Conical cap icon.svgOvate cap icon.svg Cap is conical or ovate
NA cap icon.svg Hymenium attachment is irregular or not applicable
Bare stipe icon.svg Stipe is bare
Transparent spore print icon.svg
Transparent spore print icon.svg
Spore print is cream to yellow
Mycorrhizal fungus.svgSaprotrophic fungus.svgEcology is mycorrhizal or saprotrophic
Mycomorphbox Choice.pngEdibility is choice

Morchella esculenta (commonly known as common morel, morel, yellow morel, true morel, morel mushroom, and sponge morel) is a species of fungus in the family Morchellaceae of the Ascomycota. It is one of the most readily recognized of all the edible mushrooms and highly sought after. Each fruit body begins as a tightly compressed, grayish sponge with lighter ridges, and expands to form a large yellowish sponge with large pits and ridges raised on a large white stem. The pitted yellow-brown caps measure 2–7 centimetres (1–3 inches) broad by 2–10 cm (1–4 in) tall, and are fused to the stem at its lower margin, forming a continuous hollow. The pits are rounded and irregularly arranged. The hollow stem is typically 2–9 cm (1–3+12 in) long by 2–5 cm (1–2 in) thick, and white to yellow. The fungus fruits under hardwoods and conifers during a short period in the spring, depending on the weather, and is also associated with old orchards, woods and disturbed grounds.

Description

The cap is pale brownish cream, yellow to tan or pale brown to grayish brown. The edges of the ridges are usually lighter than the pits, and somewhat oval in outline, sometimes bluntly cone-shaped with a rounded top or more elongate. Caps are hollow, attached to the stem at the lower edge, and typically about 2–7 centimetres (1–3 inches) broad by 2–10 cm (1–4 in) tall. The flesh is brittle. The stem is white to pallid or pale yellow, hollow, and straight or with a club-shaped or bulbous base. It is finely granular overall, somewhat ridged, generally about 2–9 cm (1–3+12 in) long by 2–5 cm (1–2 in) thick. [1] In age it may have brownish stains near the base. [2] It has a passing resemblance to the common stinkhorn ( Phallus impudicus ), for which it is sometimes mistaken. [3] Yellow morels are often found near wooded areas. Centipedes sometimes make their home inside these morels; infested morels usually have a hole in the top.

Microscopic characteristics

The spores range from white to cream to slightly yellow in deposit, although a spore print may be difficult to obtain given the shape of the fruit body. [4] The spores are formed in asci lining the pits—the ridges are sterile. [5] They are ellipsoidal, smooth, thin-walled, translucent (hyaline), and measure 17.5–21.9 by 8.8–11.0  µm. The asci are eight-spored, 223–300 by 19–20 µm, cylindrical, and hyaline. The paraphyses are filamentous, cylindrical, 5.8–8.8 µm wide, and hyaline. [6]

The hyphae of the stem are interwoven, hyaline, and measure 5.8–9.4 µm wide. The surface hyphae are inflated, spherical to pear-shaped, 22–44 µm wide, covered by a network of interwoven hyphae 11–16.8 µm wide with recurved cylindrical hyphal ends.

Development

Fruit bodies have successfully been grown in the laboratory. R. Ower was the first to describe the developmental stages of ascomata grown in a controlled chamber. [7] This was followed by in-depth cytological studies by Thomas Volk and Leonard (1989, 1990). To study the morel life cycle they followed the development of ascoma fruiting in association with tuberous begonias ( Begonia tuberhybrida ), from very small primordia to fully developed fruit bodies. [8] [9]

Young fruit bodies begin development in the form of a dense knot of hyphae, when suitable conditions of moisture and nutrient availability conditions have been reached. Hyphal knots are underground and cup-shaped for some time, but later emerge from the soil and develop into a stalked fruiting body. Further growth makes the hymenium convex with the asci facing towards the outer side. Because of the unequal growth of the surface of the hymenium, it becomes folded to form many ridges and depressions, resulting in the sponge or honeycomb appearance. [10]

Similar species

Gyromitra esculenta, a poisonous M. esculenta lookalike Katiuzhans'kii zakaznik Storchok zvichainii 02.jpg
Gyromitra esculenta , a poisonous M. esculenta lookalike

Morchella esculenta is probably the most familiar of the morels. In contrast to M. angusticeps and its relatives, the caps are light-colored throughout development, especially the ridges, which remain paler than the pits. M. crassipes is sometimes confused with M. esculenta. According to Smith (1975), the two are distinct, but young forms of M. crassipes are difficult to separate from M. esculenta. The two are similar in color, but M. crassipes is larger, often has thin ridges, and sometimes has a stem base that is enlarged and longitudinally grooved. [6]

Stinkhorns (esp. Phallus impudicus) have also been confused with morels, [11] but specimens of the former have a volva at the base of the stem, and are covered with gleba—a slimy, foul-smelling spore mass.

See also Gyromitra esculenta , which is similar in appearance but poisonous.

Taxonomy

The fungus was originally named Phallus esculentus by Carl Linnaeus in his Species Plantarum (1753), [12] and given its current name by Swedish mycologist Elias Magnus Fries in 1801. [13] [14]

Morchella esculenta is commonly known by various names: morel, common morel, true morel, morel mushroom, yellow morel, sponge morel, [15] Molly Moocher, haystack, and dryland fish. [2] In Nepal it is known as Guchi chyau. [16] The specific epithet is derived from the Latin esculenta, meaning "edible".

The scientific name Morchella esculenta has been applied to many similar yellow morels throughout the world. In 2014 Richard et al. used DNA analysis to restrict the M. esculenta name to a single species of yellow morel commonly found in Europe and also reported from China. [17] Other species of yellow morel, including those in North America, have received new scientific names.

Distribution and habitat

Drawing by von Albin Schmalfuss, 1897 Speisemorchel-1.jpg
Drawing by von Albin Schmalfuß, 1897

Fruit bodies are sometimes solitary, but more often in groups, on the ground in a variety of habitats. A preference for soil with a limestone base (alkaline) has been noted, [2] but they have also been found in acid soils. [18] The mushroom is usually found in early spring, in forests, orchards, yards, gardens and sometimes in recently burned areas. [1] In North America, it is sometimes referred to as the "May mushroom" due to its consistent fruiting in that month, but the time of fruiting varies locally, from February to July. It is typically the last morel species to fruit in locales where more than one species are found. [19] For example, in northern Canada and in cooler mountainous regions, morels typically do not appear until June. [20] It has been suggested that the springtime fruiting may be due to their ability to grow at low temperatures to the exclusion of competition, [21] a conclusion later corroborated by experiments correlating spore germination to soil temperatures. [22]

One author suggests the acronym PETSBASH may be used to remember the trees associated with morels: pine, elm, tulip, sassafras, beech, ash, sycamore, and hickory. [23]

In North America, it is widely distributed, but especially common in eastern North America and the Midwest. David Arora notes that "large crops can also be found around the bases of dying (but not quite dead) elms attacked by Dutch elm disease." [11] The species has been named state mushroom of Minnesota, and was the first state mushroom of any state. [24] [25]

It can also be found in Brazil and Bulgaria. [26] In Jammu and Kashmir wild mushrooms, locally known as Himalayan wild mushroom, Gucchi, Morchella conica and Morchella esculenta, are gathered and supplied as medical remedy. [27]

Cultivation

Due to the mushroom's prized fruit bodies, several attempts have been made to grow the fungus in culture. In 1901, Repin reported successfully obtaining fruit bodies in a cave in which cultures had been established in flower pots nine years previously in 1892. [28]

Mycologist Taylor Piercefield developed a method using beds of hardwood tree saplings which would be inoculated with mycelium, concentrating on the symbiotic relationship of Morchella esculenta. Later, once the mycelium network had been fully developed, the beds would be treated with potassium hydroxide to replicate the pH conditions found in soil after a forest fire. This method resulted in large, mature fruits, but was not commercially viable on a large scale. More recently, small scale commercial growers have had success growing morels by using partially shaded rows of mulched wood. The rows of mulch piles are inoculated with morel mushroom spores in a solution of water and molasses which are poured over the piles of mulch and then they are allowed to grow undisturbed for several weeks. A solution of wood ashes mixed in water and diluted is subsequently poured over the rows of wood mulch which triggers fruiting of the morels. Morels are known to appear after fires and the alkaline conditions produced by wood ash mixed with water initiate fruit body formation for most species of morels. [29]

Uses

Edibility

Harvested morels Morille.JPG
Harvested morels

Morchella esculenta, like all morels, are among the most highly prized of all edible mushrooms. [30] Raw morels have a gastrointestinal irritant, hydrazine (this has never been found in morel samples however it is assumed), but parboiling or blanching before consumption will remove it. Old fruit bodies that show signs of decay may be poisonous. [4] The mushrooms may be fried in butter or baked after being stuffed with meats and vegetables. [31] The mushrooms may also be dried by threading the caps onto string and hanging them in the sun; this process is said to concentrate the flavour. [4] One study determined the main nutritional components to be as follows (on a dry weight basis): protein 32.7%, fat 2.0%, fiber 17.6%, ash 9.7%, and carbohydrates 38.0%. [32]

In one isolated case in Germany, six people were reported to have developed neurologic effects between 6–12 hours after consumption. The effects included ataxia and visual disturbances, and lasted up to a day before disappearing without enduring effects. [33]

Bioactive compounds

Both the fruit bodies and the mycelia of M. esculenta contain an uncommon amino acid, cis-3-amino-L-proline; this amino acid does not appear to be protein bound. [34] In addition to M. esculenta, the amino acid is known to exist only in M. conica and M. crassipes. [35]

Industrial applications; solid state-fermentation

Solid-state fermentation is an industrial process to produce enzymes and to upgrade the values of existing foods, especially foods from East Asia. Solid-state fermentation is a process whereby an insoluble substrate is fermented with sufficient moisture but without free water. Solid-state fermentation, unlike that of slurry state, requires no complex fermentation controls and has many advantages over submerged liquid fermentation. M. esculenta has shown promise in degrading starch and upgrading the nutritional value of cornmeal during solid-state fermentation. [36]

M. esculenta mycelia is able to bind to and inhibit the effects of furanocoumarins, chemicals found in grapefruit that inhibit human cytochrome p450 enzymes and are responsible for the "grapefruit/drug" interaction phenomenon. [37]

Related Research Articles

<i>Morchella</i> Genus of fungi

Morchella, the true morels, is a genus of edible sac fungi closely related to anatomically simpler cup fungi in the order Pezizales. These distinctive fungi have a honeycomb appearance due to the network of ridges with pits composing their caps. Morels are prized by gourmet cooks, particularly in Catalan and French cuisine. Due to difficulties in cultivation, commercial harvesting of wild morels has become a multimillion-dollar industry in the temperate Northern Hemisphere, in particular North America, Turkey, China, the Himalayas, India, and Pakistan where these highly prized fungi are found in abundance.

<i>Spathularia flavida</i> Species of fungus

Spathularia flavida, commonly known as the yellow earth tongue, the yellow fan, or the fairy fan, is an ascomycete fungus found in coniferous forests of Asia, Europe and North America. It produces a small, fan- or spoon-shaped fruit body with a flat, wavy or lobed cream to yellow colored "head" raised on a white to cream stalk. The height is usually approximately 2–5 cm, and up to 8 cm. The fungus fruits on the ground in mosses, forest duff or humus, and fruit bodies may occur singly, in large groups, or in fairy rings. The spores produced by the fungus are needle-like, and up to 95 micrometres long. Several varieties have been described that differ largely in their microscopic characteristics. S. flavida has been described by authorities variously as inedible, of unknown edibility, or edible but tough.

<i>Gyromitra caroliniana</i> Species of fungus

Gyromitra caroliniana, known commonly as the Carolina false morel or big red, is an ascomycete fungus of the genus Gyromitra, within the Pezizales group of fungi. It is found in hardwood forests of the southeastern United States, where it fruits in early spring soon after snowmelt.

<i>Morchella elata</i> Species of fungus

Morchella elata is a species of fungus in the family Morchellaceae. It is one of many related species commonly known as black morels, and until 2012 the name M. elata was broadly applied to black morels throughout the globe. Like most members of the genus, M. elata is a popular edible fungus and is sought by many mushroom hunters.

<i>Verpa conica</i> Species of fungus

Verpa conica, commonly known as the bell morel or the early morel, is a species of fungus in the family Morchellaceae. Sometimes mistaken for a true morel, this species is an “early morel” characterized by a cap resembling a thimble that is freely attached to the stem.

<i>Verpa bohemica</i> Species of fungus in the family Morchellaceae found in northern North America, Europe, and Asia

Verpa bohemica is a species of fungus in the family Morchellaceae. Commonly known as the early morel or the wrinkled thimble-cap. The mushroom has a pale yellow or brown thimble-shaped cap—2 to 4 cm in diameter by 2 to 5 cm long—that has a surface wrinkled and ribbed with brain-like convolutions. The cap hangs from the top of a lighter-colored, brittle stem that measures up to 12 cm long by 1 to 2.5 cm thick. Microscopically, the mushroom is distinguished by its large spores, typically 60–80 by 15–18 µm, and the presence of only two spores per ascus.

<i>Amanita daucipes</i> Species of fungus

Amanita daucipes is a species of fungus in the family Amanitaceae of the mushroom order Agaricales. Found exclusively in North America, the mushroom may be recognized in the field by the medium to large white caps with pale orange tints, and the dense covering of pale orange or reddish-brown powdery conical warts on the cap surface. The mushroom also has a characteristic large bulb at the base of its stem with a blunt short rooting base, whose shape is suggestive of the common names carrot-footed lepidella, carrot-foot amanita, or turnip-foot amanita. The mushroom has a strong odor that has been described variously as "sweet and nauseous", or compared to an old ham bone, or soap. Edibility is unknown for the species, but consumption is generally not recommended due its position in the Amanita subgroup Lepidella, which contains some poisonous members.

<i>Psilocybe hispanica</i> Species of fungus in the family Hymenogastraceae

Psilocybe hispanica is a species of fungus in the family Hymenogastraceae. It produces small brown mushrooms with conical to convex caps up to 10 mm (0.4 in) in diameter and stems 16 to 25 mm long by 0.5 to 1 mm thick. Reported as new to science in 2000, it is only known from the Pyrenees mountain range in northern Spain and southwestern France, where it grows on horse dung in grass fields at elevations of 1,700 to 2,300 m. The mushroom contains the psychoactive compound psilocybin. The possible depiction of this species in the 6,000-year-old Selva Pascuala rock art suggests that it might have been used in ancient religious rituals—the oldest evidence of such usage in prehistoric Europe.

<i>Spongiforma thailandica</i> Species of fungus

Spongiforma thailandica is a species of fungus in the family Boletaceae, genus Spongiforma. The stemless sponge-like species, first described in 2009, was found in Khao Yai National Park in central Thailand, where it grows in soil in old-growth forests. The rubbery fruit body, which has a strong odor of coal-tar similar to Tricholoma sulphureum, consists of numerous internal cavities lined with spore-producing tissue. Phylogenetic analysis suggests the species is closely related to the Boletaceae genera Porphyrellus and Strobilomyces.

<i>Morchella rufobrunnea</i> Species of edible fungus in the family Morchellaceae

Morchella rufobrunnea, commonly known as the blushing morel, is a species of ascomycete fungus in the family Morchellaceae. A choice edible species, the fungus was described as new to science in 1998 by mycologists Gastón Guzmán and Fidel Tapia from collections made in Veracruz, Mexico. Its distribution was later revealed to be far more widespread after several DNA studies suggested that it is also present in the West Coast of the United States, Israel, Australia, Cyprus, Malta and Switzerland.

<i>Tylopilus tabacinus</i> Species of fungus

Tylopilus tabacinus is a species of bolete fungus in the family Boletaceae. It is characterized by a tawny-brown cap measuring up to 17.5 cm (6.9 in) in diameter, and a reticulated stem up to 16.5 cm (6.5 in) long by 6 cm (2.4 in) thick. A characteristic microscopic feature is the distinctive crystalline substance encrusted on the hyphae in the surface of the cap. The species is known from the eastern United States from Florida north to Rhode Island, and west to Mississippi, and from eastern Mexico. It is a mycorrhizal species, and associates with oak and beech trees.

<i>Morchella tridentina</i> Species of fungus

Morchella tridentina is a cosmopolitan species of ascomycete fungus in the family Morchellaceae. Commonly referred to as the mountain blond or western blond morel in North America, it produces conical, grey to buff fruit bodies that are rufescent and grow up to 20 cm (7.9 in) tall and 5 cm (2.0 in) wide. This early-diverging species is distinct within the /Elata clade due to its pale colours and has been described by many names in the past, including M. frustrata, M. quercus-ilicis, M. elatoides, M. elatoides var. elegans and M. conica var. pseudoeximia, all of which were shown to be synonyms. A widely distributed relict of the last Ice Age, M. tridentina is so far known from Argentina, Armenia, Chile, Cyprus, France, India, Israel, North America, Spain and Turkey.

<i>Morchella snyderi</i> Species of fungus

Morchella snyderi is a species of fungus in the family Morchellaceae. Described as new to science in 2012, it occurs in the montane forests of western North America, including California, Idaho, Montana, Oregon and Washington. It produces fruit bodies up to 14 cm (5.5 in) tall with ridged and pitted conical caps, and stipes that become pitted in maturity. The color of the morel is yellow to tan when young, but the cap ridges become brown to black in maturity or when dried.

<i>Morchella importuna</i> Species of fungus

Morchella importuna is a species of fungus in the family Morchellaceae described from North America in 2012. It occurs in gardens, woodchip beds, and other urban settings of northern California and the Pacific Northwest region of the United States and Canada. The fungus has also been reported from Turkey, Spain, France, Switzerland, Canada and China, although it is unknown whether this is a result of accidental introductions. It is considered a choice edible mushroom. The fruit bodies develop a distinctive ladder-like pattern of pits and ridges on the surface of their conical caps.

<i>Morchella sextelata</i> Species of fungus

Morchella sextelata is a species of ascomycete fungus in the family Morchellaceae. Described as new to science in 2012, it is found in North America. It has also been found in China, although it is not known if this is a result of an accidental introduction or natural dispersion. The fruit bodies have a roughly conical cap up to 7.5 cm (3 in) tall and 5 cm (2 in) wide, with a surface of mostly vertically arranged pits. The cap is initially yellowish to brownish, but it darkens to become almost black in maturity. The stipe is white and hollow, measuring 2–5 cm (0.8–2.0 in) high by 1–2.2 cm (0.4–0.9 in) wide.

<i>Morchella populiphila</i> Species of fungus

Morchella populiphila is a species of morel fungus native to northwestern North America. Described as new to science in 2012, its specific epithet refers to its association with black cottonwood. The morel used to be referred to as Morchella semilibera in western North American field guides until molecular analysis established that to be a strictly European species. M. populiphila occurs in California, Nevada and Oregon. Its fruit bodies grow up to 15 cm (6 in) tall with a ridged and pitted conical cap that attaches about halfway down the stipe. The cap ridges are dark brown to black in maturity, while the pits are yellowish to brownish. The fungus is edible, although not as highly valued as other morels.

<i>Morchella punctipes</i> Species of fungus

Morchella punctipes is a species of morel fungus in the family Morchellaceae. It is native to North America, found widely distributed east of the Rocky Mountains. It was first found in Michigan and described scientifically by American mycologist Charles Horton Peck in 1903.

<i>Morchella anatolica</i> Species of fungus

Morchella anatolica is a rare species of ascomycete fungus in the family Morchellaceae. It was described as new to science in 2010 from southwest Anatolia, Turkey, where it grows on moss-covered stream beds in pine forests. An ancient climatic relict, M. anatolica is restricted to the Mediterranean basin and has also been documented in Spain, Cyprus and Greece, where it is sometimes encountered with trees of the Oleaceae family. Together with its sister-species Morchella rufobrunnea, they are the earliest diverging lineages in genus Morchella, forming a distinct clade that is basal in global morel phylogenies. Because of its phylogenetic position, M. anatolica has been crucial in inferring the historical biogeography of the genus, which is estimated to have emerged somewhere in the Mediterranean region in the late Jurassic.

<i>Morchella disparilis</i> Species of fungus

Morchella disparilis is an Ascomycete fungus in the family Morchellaceae. Described as new to science in 2016, M. disparilis appears to be confined to the Mediterranean basin and is so far known from Cyprus, Greece and Spain. Its most striking feature is the exceptionally deep sinus, intermediate in depth between half-free morels of the Morchella semilibera clade and typical Distantes species.

<i>Morchella dunalii</i> Species of fungus

Morchella dunalii is a species of fungus in the family Morchellaceae (Ascomycota). A widespread species in the Mediterranean basin, M. dunalii is so far known from the Balearic islands, the islands of Corsica and Cyprus, France, Spain and Turkey, where it appears to be abundant. It fruits very early in the season on calcareous soil, usually in association with the Aleppo pine, Calabrian pine and holm oak.

References

  1. 1 2 Ammirati JF, McKenny M, Stuntz DE (1987). The New Savory Wild Mushroom. Seattle: University of Washington Press. pp. 209–10. ISBN   0-295-96480-4.
  2. 1 2 3 Roody WC. (2003). Mushrooms of West Virginia and the Central Appalachians. Lexington, Kentucky: University Press of Kentucky. p. 485. ISBN   0-8131-9039-8 . Retrieved 2010-03-22.
  3. Arora, David (1986). Mushrooms Demystified: a Comprehensive Guide to the Fleshy Fungi . Berkeley, California: Ten Speed Press. pp.  768–69. ISBN   0-89815-169-4.
  4. 1 2 3 Hall IR. (2003). Edible and Poisonous Mushrooms of the World. Portland, Or: Timber Press. pp. 239–42. ISBN   0-88192-586-1.
  5. Schalkwijk-Barendsen HME. (1991). Mushrooms of Western Canada. Edmonton: Lone Pine Publishing. pp.  381–82. ISBN   0-919433-47-2.
  6. 1 2 Ammirati J, Traquair JA, Horgen PA (1985). Poisonous Mushrooms of Canada. Fitzhenry & Whiteside in cooperation with Agriculture Canada. pp. 287–88. ISBN   978-0-88902-977-4.
  7. Ower R. (1982). "Notes on the development of the morel ascocarp: Morchella esculenta". Mycologia. 74 (1): 142–44. doi:10.2307/3792639. JSTOR   3792639.
  8. Volk TJ, Leonard TJ (1989). "Experimental studies on the morel. I. Hetrokaryon formation between mono ascosporous strains of Morchella". Mycologia. 81 (4): 523–31. doi:10.2307/3760127. JSTOR   3760127.
  9. Volk T, Leonard T (1990). "Cytology of the life-cycle of Morchella". Mycological Research. 94 (3): 399–406. doi:10.1016/S0953-7562(09)80365-1.
  10. Sharma OP. (1988). Textbook of Fungi. Boston: McGraw Hill Higher Education. pp. 193–96. ISBN   0-07-460329-9.
  11. 1 2 Arora D. (1986). Mushrooms Demystified: a Comprehensive Guide to the Fleshy Fungi. Berkeley, California: Ten Speed Press. pp. 787–88. ISBN   0-89815-169-4 . Retrieved 2010-03-22.
  12. Fries EM. (1753). Species Plantarum (in Latin). pp. 1178–79. Retrieved 2010-03-21.
  13. Persoon CH. (1801). Synopsis Methodica Fungorum. Vol. 2. p. 618. Retrieved 2010-03-21.[ permanent dead link ]
  14. Kuo M. "Morchella esculenta". MushroomExpert.Com. Retrieved 2010-03-26.
  15. Dörfelt H (2001). "Morchellaceae". In Hanelt P (ed.). Mansfeld's Encyclopedia of Agricultural and Horticultural Crops: (Except Ornamentals). Mansfeld's Encyclopedia of Agricultural and Horticultural Crops. Vol. 1. Springer. p. 17. ISBN   9783540410171 . Retrieved 2013-02-09.
  16. National register of medicinal plants. IUCN-the World Conservation Union: His Majesty's Government, Ministry of Forest and Soil Conservation. 2000. p. 61. ISBN   978-92-9144-048-1 . Retrieved 2010-03-21.
  17. Richard, Franck; Bellanger, Jean-Michel; Clowez, Philippe; Courtecuisse, Regis; Hansen, Karen; O'Donnell, Kerry; Sauve, Mathieu; Urban, Alexander; Moreau, Pierre-Arthur (30 December 2014). "True morels (Morchella, Pezizales) of Europe and North America: evolutionary relationships inferred from multilocus data and a unified taxonomy". Mycologia (Preliminary version published online). 107 (2): 359–382. doi: 10.3852/14-166 . PMID   25550303. 14-166.
  18. Metzler V, Metzler S (1992). Texas Mushrooms: a Field Guide. Austin: University of Texas Press. p. 330. ISBN   0-292-75125-7 . Retrieved 2010-03-26.
  19. McKnight VB, McKnight KH (1987). A Field Guide to Mushrooms: North America. Boston: Houghton Mifflin. pp. 322–23. ISBN   0-395-91090-0 . Retrieved 2010-03-22.
  20. Bessette A, Fischer DH (1992). Edible Wild Mushrooms of North America: a Field-to-Kitchen Guide. Austin: University of Texas Press. pp. 134–35. ISBN   0-292-72080-7 . Retrieved 2010-03-22.
  21. Baker KF, Cook RJ (1974). Biological control of plant pathogens . San Francisco: W. H. Freeman. ISBN   0-7167-0589-3.
  22. Schmidt EL. (1983). "Spore germination of and carbohydrate colonization by Morchella esculenta at different soil temperatures". Mycologia. 75 (5): 870–75. doi:10.2307/3792778. JSTOR   3792778.
  23. Rosen S. (1982). A Judge Judges Mushrooms. Highlander Pr. ISBN   0-913617-01-6.
  24. "Minnesota State Symbols: Minnesota State Mushroom". Minnesota Legislature . Retrieved June 13, 2011.
  25. "2010 Minnesota Statutes: 1.149 State Mushroom". Minnesota Office of the Revisor of Statutes. Retrieved June 13, 2011.
  26. Cortez VG, Coelho G, Guerrero RT (2004). "Morchella esculenta (Ascomycota): A rare species found in Santa Maria, Rio Grande do Sul, Brazil". Biociencias (Porto Alegre) (in Portuguese). 12 (1): 51–53.
  27. JK Medicinal Plants Introduction Centre
  28. Repin C. (1901). "Sur la culture de la Morille". Revue générale des sciences pures et appliquées. 12: 595–96. Retrieved 2010-03-21.
  29. "YouTube". YouTube Growing Morels. 2019-11-08. Archived from the original on 2021-12-21. Retrieved 2019-11-11.
  30. Phillips, Roger (2010). Mushrooms and Other Fungi of North America. Buffalo, NY: Firefly Books. p. 361. ISBN   978-1-55407-651-2.
  31. Abel D, Horn B, Kay R (1993). A Guide to Kansas Mushrooms. Lawrence: University Press of Kansas. p. 63. ISBN   0-7006-0571-1.
  32. Wahid M, Sattar A, Khan S (1988). "Composition of wild and cultivated mushrooms of Pakistan". Mushroom Journal for the Tropics. 8 (2): 47–51.
  33. Pfab R, Haberl B, Kleber J, Zilker T (2008). "Cerebellar effects after consumption of edible morels (Morchella conica, Morchella esculenta)". Clinical Toxicology. 46 (3): 259–60. doi: 10.1080/15563650701206715 . PMID   18344109.
  34. Hatanaka S-I. (1969). "A new amino acid isolated from Morchella esculenta and related species". Phytochemistry. 8 (7): 1305–08. Bibcode:1969PChem...8.1305H. doi:10.1016/S0031-9422(00)85571-5.
  35. Moriguchi M, Sada SI, Hatanaka SI (1979). "Isolation of cis-3-amino-L-proline from cultered mycelia or Morchella esculenta". Applied and Environmental Microbiology. 38 (5): 1018–19. doi:10.1128/AEM.38.5.1018-1019.1979. PMC   243624 . PMID   16345456.
  36. Zhang GP, Zhang F, Ru WM, Han JR (2009). "Solid-state fermentation of cornmeal with the ascomycete Morchella esculenta for degrading starch and upgrading nutritional value". World Journal of Microbiology and Biotechnology. 26 (1): 15–20. doi:10.1007/s11274-009-0135-y. S2CID   84930561.
  37. Myung K, Narciso JA, Manthey JA (2008). "Removal of furanocoumarins in grapefruit juice by edible fungi" (PDF). Journal of Agricultural and Food Chemistry. 56 (24): 12064–68. doi:10.1021/jf802713g. PMID   19012403. Archived from the original (PDF) on 2011-07-21. Retrieved 2010-03-21.