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Morchella

Morchella, the true morels, is a genus of edible sac fungi closely related to anatomically simpler cup fungi in the order Pezizales (division Ascomycota). 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.

Morel
A black morel in Poland
Scientific classification
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Pezizomycetes
Order: Pezizales
Family: Morchellaceae
Genus: Morchella
Dill. ex Pers. : Fr. (1794)
Type species
Morchella esculenta
(L.) Pers. : Fr. (1801)
Species

~70 worldwide (see text)

Synonyms[1]
  • Phalloboletus Adans. (1763)
  • Boletus Tourn. ex Adans. (1763)
  • Eromitra Lév. (1846)
  • Mitrophora Lév. (1846)
  • Morilla Quél. (1886)
  • Morchella sect. Mitrophorae (Lév.) S.Imai (1932)

Typified by Morchella esculenta in 1794, the genus has been the source of considerable taxonomical controversy throughout the years, mostly with regard to the number of species involved, with some mycologists recognising as few as three species and others over thirty. Current molecular phylogenetics suggest there might be over seventy species of Morchella worldwide, most of them exhibiting high continental endemism and provincialism.

The genus is currently the focus of extensive phylogenetic, biogeographical, taxonomical and nomenclatural studies, and several new species have been described from Australia, Canada, Cyprus, Israel, Spain, and Turkey.

Description edit

Morels resemble a honeycomb due to the network of ridges with pits composing their caps.

Similar species edit

 
Gyromitra esculenta, a false morel

When gathering morels for the table, care must be taken to distinguish them from the poisonous "false morels", a term loosely applied to describe Gyromitra esculenta, Verpa bohemica, and other morel lookalikes. Although false morels are sometimes eaten without ill effect, they can cause severe gastrointestinal upset, loss of muscular coordination (including cardiac muscle), or even death.[2][3] Incidents of poisoning usually occur when they are eaten in large quantities, inadequately cooked, or over several days in a row. False morels contain gyromitrin, an organic carcinogenic poison, hydrolyzed in the body into monomethylhydrazine (MMH).[4] Gyromitra esculenta in particular, has been reported to be responsible for up to 23% of mushroom fatalities each year in Poland.[5]

The key morphological features distinguishing false morels from true morels are as follows:

  • Gyromitra species often have a "wrinkled" or "cerebral" (brain-like) appearance to the cap due to multiple wrinkles and folds, rather than the honeycomb appearance of true morels due to ridges and pits.
  • Gyromitra esculenta has a cap that is usually reddish-brown in colour, but sometimes also chestnut, purplish-brown, or dark brown.
  • Gyromitra species are typically chambered in longitudinal section, while Verpa species contain a cottony substance inside their stem, in contrast to true morels which are always hollow.
  • The caps of Verpa species (V. bohemica, V. conica and others) are attached to the stem only at the apex (top of the cap), unlike true morels which have caps that are attached to the stem at, or near the base of the cap. The easiest way to distinguish Verpa species from Morchella species is to slice them longitudinally.[6][7]

Taxonomy edit

The fruit bodies of Morchella species are highly polymorphic, varying in shape, color, and size. While in many cases they do not exhibit clear-cut distinguishing features microscopically, this has historically contributed to uncertainties in taxonomy.[8][9][10][11] Discriminating between the various taxa described is further hindered by uncertainty over which of these are truly biologically distinct. Remarkably, some authors in the past had suggested that the genus contains as few as 3 to 6 species,[12][13][14][15][16] while others recognised as many as 34.[17][18][19] Efforts to clarify the situation and re-evaluate old classical names (such as Morchella elata and others) in accordance to current phylogenetic data have been challenging, due to vague or ambiguous original descriptions and loss of holotype material.[20][21] In 2012, the simultaneous description of several new taxa from Europe by Clowez[19] and North America by Kuo and colleagues[22] resulted in several synonymities further complicating matters, until a transatlantic study by Richard and colleagues resolved many of these issues in 2014.[20] The genus is currently undergoing extensive re-evaluation with regards to the taxonomic status of several species.

Early taxonomic history edit

Morchella Dill. ex Pers. : Fr. was typified by Christiaan Hendrik Persoon in 1794,[23] with Morchella esculenta designated as the type species for the genus. Among early pioneers who took an interest in the genus, were mycologists Julius Vincenz von Krombholz and Émile Boudier, who, in 1834[24] and 1897[17] respectively, published several species and varieties, accompanied by meticulously illustrated iconographic plates. The seminal taxon Morchella elata, whose true identity still remains unresolved,[20][11] was described by Elias Fries in 1822, from a fir forest in Sweden.[25] Other classical, early-proposed names include Morchella deliciosa, also described by Fries in 1822, Morchella semilibera, the half-free morel, originally described by de Candolle and sanctioned by Fries in 1822,[25] Morchella vulgaris, which was recombined by Samuel Gray as a distinct species in 1821[26] following a forma of M. esculenta previously proposed by Persoon, and Morchella angusticeps, a large-spored species described by American mycologist Charles Peck in 1887.[27] Morchella purpurascens, the purple morel, was first described by Boudier as a variety of M. elata in 1897 based on an 1834 plate by Krombholz, and was recombined as a distinct species in 1985 by Emile Jacquetant.[18][28] Morchella eximia, a globally-occurring fire-associated species was also described by Boudier in 1910.[29] The old, widely applied name Morchella conica,[30] featuring in many field guides and literature across several countries, has been shown by Richard and colleagues to be illegitimate.[20]

Classification edit

About 80 species of Morchella were described until the turn of the 21st century (http://www.indexfungorum.org/), a number of which were later shown to be illegitimate or synonyms.[20] As molecular tools became widely available in the new millennium, a revived interest in the genus commenced and several new species were proposed. In 2008 Kuo described Morchella tomentosa from burned coniferous forests in western North America.[31] In 2010 Işiloğlu and colleagues described Morchella anatolica,[32] a basal species from Turkey later shown to be sister to Morchella rufobrunnea. A study by Clowez described over 20 new species in 2012,[19] while later in the same year, another study by Kuo and colleagues described 19 species from North America.[22] However, several of these newly proposed names later turned out to be synonyms.[20] An extensive taxonomical and nomenclatural revision of the genus provided by Richard and colleagues in 2014, applied names to 30 of the genealogical lineages recognized so far and clarified several synonymities.[20] Also in 2014, Elliott and colleagues described Morchella australiana from sclerophyll forests in Australia,[33] while Clowez and colleagues described Morchella fluvialis from riparian forests in Spain.[34]

In 2015, Loizides and colleagues clarified the taxonomy of Morchella tridentina, a cosmopolitan species described under many names, and recombined Morchella kakiicolor as a distinct species.[21] Later in the same year, Clowez and colleagues described Morchella palazonii from Spain,[35] while Voitk and colleagues described Morchella laurentiana from Canada and Morchella eohespera, a cosmopolitan species present in several continents.[36] In an extensive phylogenetic and morphological study from Cyprus in 2016, Loizides and colleagues added two more Mediterranean species, Morchella arbutiphila and Morchella disparilis, and resurrected Morchella dunensis as an autonomous species.[37] In the same year, Taşkın and colleagues described four of the previously unnamed phylospecies from Turkey: Morchella conifericola, Morchella feekensis, Morchella magnispora and Morchella mediteterraneensis.[38]

Section Rufobrunnea edit

Section Morchella edit

Section Distantes edit

Unresolved classification edit

Phylogeny edit

Early phylogenetic analyses supported the hypothesis that the genus comprises only a few species with considerable phenotypic variation.[40][41][42] Subsequent multigenic DNA studies, however, have revealed more than a dozen genealogically distinct species in North America and at least as many in Europe.[43][44][45][20] DNA studies revealed three discrete clades, or genetic groups, consisting of the "white morels" (Morchella rufobrunnea and Morchella anatolica), the "yellow morels" (Morchella esculenta and others), and the "black morels" (Morchella elata and others).[44] The fire-associated species Morchella tomentosa, commonly known as the "gray morel", is distinct for its fine hairs on the cap ridges and sclerotia-like underground structures, and may also deserve its own clade based on DNA evidence.[46][31][47] Within the yellow and black clades, there are dozens of distinct species, many endemic to individual continents or regions.[44] This species-rich view is supported by studies in Western Europe,[48] Turkey,[49] Cyprus,[37] Israel,[50] China,[51] Patagonia,[52] and the Himalayas.[53]

Early ancestral reconstruction tests by O'Donnell and collaborators postulated a western North American origin of morels and the genus was estimated to have diverged from its closest genealogical relatives Verpa and Disciotis in the early Cretaceous, approximately 129 million years ago (Mya).[44] This date was later revised by Du and collaborators, placing the divergence of the genus in the late Jurassic, approximately 154 Mya.[51] However, neither of these reconstructions had included Morchella anatolica in the analyses, whose phylogenetic placement remained at the time unresolved. Following genetic testing of isotype collection of M. anatolica by Taşkın and colleagues, this species was shown to nest in the ancestral /Rufobrunnea clade, together with the transcontinental Morchella rufobrunnea.[45] This cast doubts over the accuracy of the original reconstructions, since both species of the ancestral /Rufobrunnea clade are present in the Mediterranean, while M. anatolica is altogether absent from North America.[21][37] Updated ancestral area reconstructions by Loizides and colleagues using an expanded 79-species data set, have in 2021 refuted the previous hypothesis and designated the Mediterranean basin as the most probable place of origin of morels.[54]

Distribution and ecology edit

 
Yellow morels in West Virginia, USA

The ecology of Morchella species is not well understood. Many species appear to form symbiotic or endophytic relationships with trees,[55][56][57][58] while others appear to act as saprotrophs.[46][57] Yellow morels (Morchella esculenta and related species) are more commonly found under deciduous trees rather than conifers, while black morels (Morchella elata and related species) are mostly found in coniferous forests, disturbed ground and recently burned areas.[19][37][59][60] Morchella galilaea,[61] and occasionally Morchella rufobrunnea,[50][62][21] appear to fruit in the autumn or winter months rather than spring, which is the typical fruiting season for morels.

Tree species associated with Morchella vary greatly depending on the individual species, continent, or region. Trees commonly associated with morels in Europe and across the Mediterranean include Abies (fir), Pinus (pine), Populus (poplar), Ulmus (elm), Quercus (oak), Arbutus (strawberry trees), Castanea (chestnut), Alnus (alder), Olea (olive trees), Malus (apple trees), and Fraxinus (ash).[19][37][21][45][54] In western North America morels are often found in coniferous forests, including species of Pinus (pine), Abies (fir), Larix (larch), and Pseudotsuga (Douglas-fir), as well as in Populus (cottonwood) riparian forests.[63][22] Deciduous trees commonly associated with morels in the northern hemisphere include Fraxinus (ash), Platanus (sycamore), Liriodendron (tulip tree), dead and dying elms, cottonwoods, and old apple trees (remnants of orchards).[22] Due to their springtime phenology (March–May), morels are hardly ever found in the vicinity of common poisonous mushrooms such as the death cap (Amanita phalloides), the sulphur tuft (Hypholoma fasciculare), or the fly agaric (Amanita muscaria).[64] They can, however, occur alongside false morels (Gyromitra and Verpa species) and elfin saddles (Helvella species), which also appear in spring.

Efforts to cultivate morels at a large scale have rarely been successful and the commercial morel industry relies on the harvest of wild mushrooms.[63]

Association with wildfire edit

 
Longneck morel in Indiana, USA

Certain Morchella species (M. eximia, M. importuna, M. tomentosa and others) exhibit a pyrophilic behaviour and may grow abundantly in forests which have been recently burned by a fire.[65][66] Moderate-intensity fires are reported to produce higher abundances of morels than low- or high-intensity fires.[47] This is caused by the soil becoming more alkaline as the result of wood ash combining with water and being absorbed into the soil which triggers the morels to fruit. Alkaline soil conditions which trigger fruiting have been observed and exploited with small-scale commercial cultivation of morels.[67][46][65] Where fire suppression is practiced, morels often grow in small numbers in the same spot, year after year. If these areas are overrun by wildfire they often produce a bumper crop of black morels the following spring. Commercial pickers and buyers in North America target recently burned areas for this reason. These spots may be closely guarded by mushroom pickers, as morels are widely regarded as a delicacy and often a cash crop.[63]

Transcontinental species edit

 
Black morel in Washington (state), USA

Although many species within Morchella exhibit continental endemism and provincialism,[44] several species have been phylogenetically shown to be present in more than one continent. So far, the list of transcontinental species includes M. americana, M. eohespera, M. eximia, M. exuberans, M. galilaea, M. importuna, M. populiphila, M. pulchella, M. rufobrunnea, M. semilibera, M. sextelata, M. steppicola, and M. tridentina.[51][20][37][11] The reasons behind the widespread, cosmopolitan distribution of these species, are still puzzling. Some authors have hypothesized that such transcontinental occurrences are the result of accidental anthropogenic introductions,[49][44] but this view has been disputed by others, who suggested an old and natural distribution, at least for some of these species which appear to be linked to indigenous flora.[21][37][54] Long-distance spore dispersal has also been suggested as a possible dispersal mechanism for some species, especially those belonging to fire-adapted lineages.[68] It has been suggested that the widespread but disjunct distribution of some morel species, especially early diverging lineages like M. rufobrunnea and M. tridentina, may be the result of climatic refugia from the Quaternary glaciation.[54]

Cultivation edit

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.[69]

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 alkalinity produced by wood ash mixed with water initiate fruit body formation for most species of morels.[67]

In 2021 it was announced that indoor cultivation of black morels had been successfully achieved after decades of research and experimentation with methods by The Danish Morel Project. The project has been able to cultivate 20 lbs of morels per square yard or around 10 kg per square metre with cost estimates expected to be similar to producing white button mushrooms (Agaricus bisporus). Previous attempts at cultivation had managed to produce sclerotia but encountered issues in getting them to reliably fruit. One of the breakthroughs with this project was growing them in a climate controlled environment in conjunction with grass which is involved in stimulating fruiting in the morel mycelium. Cultivation in this manner has been noted to produce superior morels for culinary uses since they can be assured to be insect, slug and dirt free and therefore do not need to be washed and cleaned like foraged morels. Since washing morels can negatively impact the texture, reliable cultivation may result in more versatility with this ingredient in the kitchen as well as making the delicacy more affordable and accessible.[70][71]

Toxicity edit

Morchella species are thought to contain small amounts of hydrazine[72] toxins or an unknown toxin that is destroyed through cooking (the presence of hydrazine is controversial since there are no primary references of hydrazine having been detected in the species); because of this, morels should never be eaten raw.[73] It has been reported that even cooked morels can sometimes cause symptoms of upset stomach when consumed with alcohol.[74]

When eating this fungus for the first time it is wise to consume a small amount to minimize any allergic reaction. As with all fungi, morels for consumption must be clean and free of decay. Morels growing in old apple orchards previously treated with the now-banned insecticide lead arsenate may accumulate levels of toxic lead and arsenic that are unsuitable for human consumption.[75]

Uses edit

Morels, "almost universally associated with spring," can be found in many habitats. Morel may be more likely to fruit during a period of increasing heat following a chilly period, a preference which is credited for their abundance in areas with cold winters.[76]

Black morels (Morchella elata) are often found on land that has been disturbed by logging burning.[76]

Nutrition edit

Morel mushrooms, raw
Nutritional value per 100 g (3.5 oz)
Energy129 kJ (31 kcal)
5.1 g
Sugars0.6 g
Dietary fiber2.8 g
0.57 g
3.12 g
VitaminsQuantity
%DV
Thiamine (B1)
6%
0.069 mg
Riboflavin (B2)
17%
0.205 mg
Niacin (B3)
15%
2.252 mg
Pantothenic acid (B5)
9%
0.44 mg
Vitamin B6
10%
0.136 mg
Folate (B9)
2%
9 μg
Vitamin D
34%
5.1 μg
MineralsQuantity
%DV
Calcium
4%
43 mg
Iron
94%
12.18 mg
Magnesium
5%
19 mg
Manganese
28%
0.587 mg
Phosphorus
28%
194 mg
Potassium
14%
411 mg
Zinc
21%
2.03 mg
Other constituentsQuantity
Water90 g

Link to USDA Database entry
Percentages are roughly approximated using US recommendations for adults.
Source: USDA FoodData Central

Raw morel mushrooms are 90% water, 5% carbohydrates, 3% protein, and 1% fat. A 100 gram reference amount supplies 31 calories, and is a rich source of iron (94% of the Daily Value, DV), manganese, phosphorus, zinc, and vitamin D (34% DV, if having been exposed to sunlight or artificial ultraviolet light). Raw morels contain moderate levels of several B vitamins (table).

Gastronomical value and culinary uses edit

They have been called "prized delicacies...they are so esteemed in Europe that people used to set fire to their own forests in hopes of eliciting a bountiful morel crop the next spring!"[76]

Morels are a feature of many cuisines, including Provençal.[77] Their flavor is prized by chefs worldwide, with recipes and preparation methods designed to highlight and preserve it.[78] As with most edible fungi, they are best when collected or bought fresh. They are sometimes added to meat and poultry dishes and soups, and can be used as pasta fillings.[79] As morels are known to contain thermolabile toxins, they must always be cooked before eating.

Morels can be preserved in several ways: They can be 'flash frozen' by simply running under cold water or putting them in a bucket to soak for a few minutes, then spread on a baking tray and placed into a freezer. After freezing, they keep very well with the frozen glaze for a long time in airtight containers. However, when thawed they can sometimes turn slightly mushy, so they are best frozen after steaming or frying. Due to their natural porosity, morels may contain trace amounts of soil which cannot be easily washed out. Any visible soil should be removed with a brush, after cutting the body in half lengthwise, if needed. Mushroom hunters sometimes recommend soaking morels in a bowl of salt water briefly prior to cooking, although many chefs would disagree.[80]

Drying is a popular and effective method for long-term storage, and morels are widely available commercially in this form. Any insect larvae which might be present in the fruit bodies usually drop out during the drying process.[81] Dried morels can then be reconstituted by soaking for 10–20 minutes in warm water or milk, and the soaking liquid can be used as stock.[82]

The flavor of morels is not just appreciated by humans; in Yellowstone National Park, black morels are also known to be consumed by grizzly bears (Ursus arctos horribilis).[83]

In popular culture edit

Morel hunting is a common springtime activity. Mushroom collectors may carry a mesh collecting bag, so the spores can scatter as one carries the harvest.[80]

Every spring, hundreds of morel enthusiasts gather in Boyne City, Michigan for the National Morel Mushroom Festival, a century-old event.[84] As one observer stated, "if there is a modern, North American reenactment of Geoffrey Chaucer's Canterbury Tales this is it."[7] Other festivals and hunting competitions in North America include the Illinois State Morel Mushroom Hunting Championship, the Ottawa Midwest Morel Fest and the Mesick Michigan Mushroom Festival.[85]

In the survival horror video game Cataclysm: Dark Days Ahead, morel mushrooms are featured as one of the many edible items that the survivors can gather in the forests. They typically can be picked from underbrushes during summer and autumn, and can be boiled, fried, or preserved via drying.

In the farming sim video game Stardew Valley, morels are a consumable item that can be found in the Secret Woods or Forest Farm during spring. Its player chooses to allow mushrooms to grow in the Farm Cave, morels have an 8.1% chance of spawning each day regardless of season. The item's description reads that it is "sought after for its unique and nutty flavor".

Vernacular names edit

Morchella species have been called by many local names; some of the more colorful include dryland fish, because when sliced lengthwise then breaded and fried, their outline resembles the shape of a fish;[86] hickory chickens, as they are known in many parts of Kentucky; and merkels or miracles, based on folklore, of how a mountain family was saved from starvation by eating morels. In parts of West Virginia, they are known as molly moochers, muggins, or muggles. Due to the partial structural and textural similarity to some species of Porifera (sponges), other common names for any true morel are sponge mushroom and waffle mushroom. In the Appalachian woodlands, morels have also been called haystacks, or snakeheads.[80] The Finnish vernacular name huhtasieni, refers to huhta, area cleared for agriculture by the slash and burn method.[87]

The scientific name of the genus Morchella itself, is thought to have derived from morchel,[88] an old German word close to "Möhre", carrot or beet, due to similarity in shape.

See also edit

References edit

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External links edit

  • 2000 Pezizales on Ascomycete.org
  • MushroomExpert.com's Morel section
  • "A Beginner's Guide to Hunting Morel Mushrooms", from Field and Stream
  • Colorado Morels 2020-11-11 at the Wayback Machine

morchella, morel, redirects, here, other, uses, morel, disambiguation, true, morels, genus, edible, fungi, closely, related, anatomically, simpler, fungi, order, pezizales, division, ascomycota, these, distinctive, fungi, have, honeycomb, appearance, network, . Morel redirects here For other uses see Morel disambiguation Morchella the true morels is a genus of edible sac fungi closely related to anatomically simpler cup fungi in the order Pezizales division Ascomycota 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 MorelA black morel in PolandScientific classificationDomain EukaryotaKingdom FungiDivision AscomycotaClass PezizomycetesOrder PezizalesFamily MorchellaceaeGenus MorchellaDill ex Pers Fr 1794 Type speciesMorchella esculenta L Pers Fr 1801 Species 70 worldwide see text Synonyms 1 Phalloboletus Adans 1763 Boletus Tourn ex Adans 1763 Eromitra Lev 1846 Mitrophora Lev 1846 Morilla Quel 1886 Morchella sect Mitrophorae Lev S Imai 1932 Typified by Morchella esculenta in 1794 the genus has been the source of considerable taxonomical controversy throughout the years mostly with regard to the number of species involved with some mycologists recognising as few as three species and others over thirty Current molecular phylogenetics suggest there might be over seventy species of Morchella worldwide most of them exhibiting high continental endemism and provincialism The genus is currently the focus of extensive phylogenetic biogeographical taxonomical and nomenclatural studies and several new species have been described from Australia Canada Cyprus Israel Spain and Turkey Contents 1 Description 1 1 Similar species 2 Taxonomy 2 1 Early taxonomic history 2 2 Classification 2 2 1 Section Rufobrunnea 2 2 2 Section Morchella 2 2 3 Section Distantes 2 2 4 Unresolved classification 2 3 Phylogeny 3 Distribution and ecology 3 1 Association with wildfire 3 2 Transcontinental species 4 Cultivation 5 Toxicity 6 Uses 6 1 Nutrition 6 2 Gastronomical value and culinary uses 7 In popular culture 7 1 Vernacular names 8 See also 9 References 10 External linksDescription editThis section needs expansion You can help by adding to it November 2023 Morels resemble a honeycomb due to the network of ridges with pits composing their caps nbsp nbsp nbsp nbsp nbsp nbsp Similar species edit nbsp Gyromitra esculenta a false morelWhen gathering morels for the table care must be taken to distinguish them from the poisonous false morels a term loosely applied to describe Gyromitra esculenta Verpa bohemica and other morel lookalikes Although false morels are sometimes eaten without ill effect they can cause severe gastrointestinal upset loss of muscular coordination including cardiac muscle or even death 2 3 Incidents of poisoning usually occur when they are eaten in large quantities inadequately cooked or over several days in a row False morels contain gyromitrin an organic carcinogenic poison hydrolyzed in the body into monomethylhydrazine MMH 4 Gyromitra esculenta in particular has been reported to be responsible for up to 23 of mushroom fatalities each year in Poland 5 The key morphological features distinguishing false morels from true morels are as follows Gyromitra species often have a wrinkled or cerebral brain like appearance to the cap due to multiple wrinkles and folds rather than the honeycomb appearance of true morels due to ridges and pits Gyromitra esculenta has a cap that is usually reddish brown in colour but sometimes also chestnut purplish brown or dark brown Gyromitra species are typically chambered in longitudinal section while Verpa species contain a cottony substance inside their stem in contrast to true morels which are always hollow The caps of Verpa species V bohemica V conica and others are attached to the stem only at the apex top of the cap unlike true morels which have caps that are attached to the stem at or near the base of the cap The easiest way to distinguish Verpa species from Morchella species is to slice them longitudinally 6 7 Taxonomy editThe fruit bodies of Morchella species are highly polymorphic varying in shape color and size While in many cases they do not exhibit clear cut distinguishing features microscopically this has historically contributed to uncertainties in taxonomy 8 9 10 11 Discriminating between the various taxa described is further hindered by uncertainty over which of these are truly biologically distinct Remarkably some authors in the past had suggested that the genus contains as few as 3 to 6 species 12 13 14 15 16 while others recognised as many as 34 17 18 19 Efforts to clarify the situation and re evaluate old classical names such as Morchella elata and others in accordance to current phylogenetic data have been challenging due to vague or ambiguous original descriptions and loss of holotype material 20 21 In 2012 the simultaneous description of several new taxa from Europe by Clowez 19 and North America by Kuo and colleagues 22 resulted in several synonymities further complicating matters until a transatlantic study by Richard and colleagues resolved many of these issues in 2014 20 The genus is currently undergoing extensive re evaluation with regards to the taxonomic status of several species Early taxonomic history edit Morchella Dill ex Pers Fr was typified by Christiaan Hendrik Persoon in 1794 23 with Morchella esculenta designated as the type species for the genus Among early pioneers who took an interest in the genus were mycologists Julius Vincenz von Krombholz and Emile Boudier who in 1834 24 and 1897 17 respectively published several species and varieties accompanied by meticulously illustrated iconographic plates The seminal taxon Morchella elata whose true identity still remains unresolved 20 11 was described by Elias Fries in 1822 from a fir forest in Sweden 25 Other classical early proposed names include Morchella deliciosa also described by Fries in 1822 Morchella semilibera the half free morel originally described by de Candolle and sanctioned by Fries in 1822 25 Morchella vulgaris which was recombined by Samuel Gray as a distinct species in 1821 26 following a forma of M esculenta previously proposed by Persoon and Morchella angusticeps a large spored species described by American mycologist Charles Peck in 1887 27 Morchella purpurascens the purple morel was first described by Boudier as a variety of M elata in 1897 based on an 1834 plate by Krombholz and was recombined as a distinct species in 1985 by Emile Jacquetant 18 28 Morchella eximia a globally occurring fire associated species was also described by Boudier in 1910 29 The old widely applied name Morchella conica 30 featuring in many field guides and literature across several countries has been shown by Richard and colleagues to be illegitimate 20 Classification edit About 80 species of Morchella were described until the turn of the 21st century http www indexfungorum org a number of which were later shown to be illegitimate or synonyms 20 As molecular tools became widely available in the new millennium a revived interest in the genus commenced and several new species were proposed In 2008 Kuo described Morchella tomentosa from burned coniferous forests in western North America 31 In 2010 Isiloglu and colleagues described Morchella anatolica 32 a basal species from Turkey later shown to be sister to Morchella rufobrunnea A study by Clowez described over 20 new species in 2012 19 while later in the same year another study by Kuo and colleagues described 19 species from North America 22 However several of these newly proposed names later turned out to be synonyms 20 An extensive taxonomical and nomenclatural revision of the genus provided by Richard and colleagues in 2014 applied names to 30 of the genealogical lineages recognized so far and clarified several synonymities 20 Also in 2014 Elliott and colleagues described Morchella australiana from sclerophyll forests in Australia 33 while Clowez and colleagues described Morchella fluvialis from riparian forests in Spain 34 In 2015 Loizides and colleagues clarified the taxonomy of Morchella tridentina a cosmopolitan species described under many names and recombined Morchella kakiicolor as a distinct species 21 Later in the same year Clowez and colleagues described Morchella palazonii from Spain 35 while Voitk and colleagues described Morchella laurentiana from Canada and Morchella eohespera a cosmopolitan species present in several continents 36 In an extensive phylogenetic and morphological study from Cyprus in 2016 Loizides and colleagues added two more Mediterranean species Morchella arbutiphila and Morchella disparilis and resurrected Morchella dunensis as an autonomous species 37 In the same year Taskin and colleagues described four of the previously unnamed phylospecies from Turkey Morchella conifericola Morchella feekensis Morchella magnispora and Morchella mediteterraneensis 38 Section Rufobrunnea edit Morchella anatolica synonym Morchella lanceolata Morchella rufobrunneaSection Morchella edit Morchella americana synonyms Morchella californica Morchella claviformis Morchella esculentoides Morchella populina Morchella castaneae synonyms Morchella brunneorosea Morchella brunneorosea var sordida Morchella diminutiva Morchella dunensis synonyms Morchella esculenta f dunensis Morchella andalusiae Morchella esculenta synonyms Morchella pseudoumbrina Morchella pseudoviridis Morchella fluvialis Morchella galilaea Morchella palazonii Morchella prava Morchella sceptriformis synonym Morchella virginiana Morchella steppicola Morchella ulmaria synonym Morchella cryptica Morchella vulgaris synonyms Morchella acerina Morchella anthracina Morchella lepida Morchella robiniae Morchella spongiolaSection Distantes edit Morchella angusticeps Morchella arbutiphila Morchella australiana Morchella brunnea Morchella conifericola Morchella deliciosa synonym Morchella conica Morchella disparilis Morchella dunalii synonym Morchella fallax Morchella elata Morchella eohespera Morchella eximia synonyms Morchella anthracophila Morchella carbonaria Morchella septimelata Morchella eximioides Morchella exuberans synonym Morchella capitata Morchella feekensis Morchella iberica 39 Morchella importuna Morchella kakiicolor synonym Morchella quercus ilicis f kakiicolor Morchella laurentiana Morchella magnispora Morchella mediteterraneensis Morchella populiphila Morchella pulchella Morchella punctipes Morchella purpurascens synonyms Morchella elata var purpurascens Morchella conica Morchella conica var purpurascens Morchella conica var crassa Morchella semilibera synonyms Morchella gigas Morchella gigas var tintinnabulum Morchella hybrida Morchella undosa Morchella varisiensis Morchella esculenta var crassipes Phallus gigas Eromitra gigas Phallus undosus Phallus crassipes Mitrophora hybrida Mitrophora hybrida var crassipes Ptychoverpa gigas Helvella hybrida Morchella septentrionalis Morchella sextelata Morchella snyderi Morchella tomentosa Morchella tridentina synonyms Morchella quercus ilicis Morchella frustrata Morchella elatoides Morchella elatoides var elagans Morchella conica var pseudoeximiaUnresolved classification edit Morchella anteridiformis Morchella apicata Morchella bicostata Morchella conicopapyracea Morchella crassipes Morchella deqinensis Morchella distans Morchella guatemalensis Morchella herediana Morchella hetieri Morchella hortensis Morchella hotsonii Morchella hungarica Morchella inamoena Morchella intermedia Morchella meiliensis Morchella miyabeana Morchella neuwirthii Morchella norvegiensis Morchella patagonica Morchella patula Morchella pragensis Morchella procera Morchella pseudovulgaris Morchella rielana Morchella rigida Morchella rigidoides Morchella smithiana Morchella sulcata Morchella tasmanica Morchella tatari Morchella tibetica Morchella umbrina Morchella umbrinovelutipes Morchella vaporaria Phylogeny edit Early phylogenetic analyses supported the hypothesis that the genus comprises only a few species with considerable phenotypic variation 40 41 42 Subsequent multigenic DNA studies however have revealed more than a dozen genealogically distinct species in North America and at least as many in Europe 43 44 45 20 DNA studies revealed three discrete clades or genetic groups consisting of the white morels Morchella rufobrunnea and Morchella anatolica the yellow morels Morchella esculenta and others and the black morels Morchella elata and others 44 The fire associated species Morchella tomentosa commonly known as the gray morel is distinct for its fine hairs on the cap ridges and sclerotia like underground structures and may also deserve its own clade based on DNA evidence 46 31 47 Within the yellow and black clades there are dozens of distinct species many endemic to individual continents or regions 44 This species rich view is supported by studies in Western Europe 48 Turkey 49 Cyprus 37 Israel 50 China 51 Patagonia 52 and the Himalayas 53 Early ancestral reconstruction tests by O Donnell and collaborators postulated a western North American origin of morels and the genus was estimated to have diverged from its closest genealogical relatives Verpa and Disciotis in the early Cretaceous approximately 129 million years ago Mya 44 This date was later revised by Du and collaborators placing the divergence of the genus in the late Jurassic approximately 154 Mya 51 However neither of these reconstructions had included Morchella anatolica in the analyses whose phylogenetic placement remained at the time unresolved Following genetic testing of isotype collection of M anatolica by Taskin and colleagues this species was shown to nest in the ancestral Rufobrunnea clade together with the transcontinental Morchella rufobrunnea 45 This cast doubts over the accuracy of the original reconstructions since both species of the ancestral Rufobrunnea clade are present in the Mediterranean while M anatolica is altogether absent from North America 21 37 Updated ancestral area reconstructions by Loizides and colleagues using an expanded 79 species data set have in 2021 refuted the previous hypothesis and designated the Mediterranean basin as the most probable place of origin of morels 54 Distribution and ecology edit nbsp Yellow morels in West Virginia USAThe ecology of Morchella species is not well understood Many species appear to form symbiotic or endophytic relationships with trees 55 56 57 58 while others appear to act as saprotrophs 46 57 Yellow morels Morchella esculenta and related species are more commonly found under deciduous trees rather than conifers while black morels Morchella elata and related species are mostly found in coniferous forests disturbed ground and recently burned areas 19 37 59 60 Morchella galilaea 61 and occasionally Morchella rufobrunnea 50 62 21 appear to fruit in the autumn or winter months rather than spring which is the typical fruiting season for morels Tree species associated with Morchella vary greatly depending on the individual species continent or region Trees commonly associated with morels in Europe and across the Mediterranean include Abies fir Pinus pine Populus poplar Ulmus elm Quercus oak Arbutus strawberry trees Castanea chestnut Alnus alder Olea olive trees Malus apple trees and Fraxinus ash 19 37 21 45 54 In western North America morels are often found in coniferous forests including species of Pinus pine Abies fir Larix larch and Pseudotsuga Douglas fir as well as in Populus cottonwood riparian forests 63 22 Deciduous trees commonly associated with morels in the northern hemisphere include Fraxinus ash Platanus sycamore Liriodendron tulip tree dead and dying elms cottonwoods and old apple trees remnants of orchards 22 Due to their springtime phenology March May morels are hardly ever found in the vicinity of common poisonous mushrooms such as the death cap Amanita phalloides the sulphur tuft Hypholoma fasciculare or the fly agaric Amanita muscaria 64 They can however occur alongside false morels Gyromitra and Verpa species and elfin saddles Helvella species which also appear in spring Efforts to cultivate morels at a large scale have rarely been successful and the commercial morel industry relies on the harvest of wild mushrooms 63 Association with wildfire edit nbsp Longneck morel in Indiana USACertain Morchella species M eximia M importuna M tomentosa and others exhibit a pyrophilic behaviour and may grow abundantly in forests which have been recently burned by a fire 65 66 Moderate intensity fires are reported to produce higher abundances of morels than low or high intensity fires 47 This is caused by the soil becoming more alkaline as the result of wood ash combining with water and being absorbed into the soil which triggers the morels to fruit Alkaline soil conditions which trigger fruiting have been observed and exploited with small scale commercial cultivation of morels 67 46 65 Where fire suppression is practiced morels often grow in small numbers in the same spot year after year If these areas are overrun by wildfire they often produce a bumper crop of black morels the following spring Commercial pickers and buyers in North America target recently burned areas for this reason These spots may be closely guarded by mushroom pickers as morels are widely regarded as a delicacy and often a cash crop 63 Transcontinental species edit nbsp Black morel in Washington state USAAlthough many species within Morchella exhibit continental endemism and provincialism 44 several species have been phylogenetically shown to be present in more than one continent So far the list of transcontinental species includes M americana M eohespera M eximia M exuberans M galilaea M importuna M populiphila M pulchella M rufobrunnea M semilibera M sextelata M steppicola and M tridentina 51 20 37 11 The reasons behind the widespread cosmopolitan distribution of these species are still puzzling Some authors have hypothesized that such transcontinental occurrences are the result of accidental anthropogenic introductions 49 44 but this view has been disputed by others who suggested an old and natural distribution at least for some of these species which appear to be linked to indigenous flora 21 37 54 Long distance spore dispersal has also been suggested as a possible dispersal mechanism for some species especially those belonging to fire adapted lineages 68 It has been suggested that the widespread but disjunct distribution of some morel species especially early diverging lineages like M rufobrunnea and M tridentina may be the result of climatic refugia from the Quaternary glaciation 54 Cultivation editDue 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 69 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 alkalinity produced by wood ash mixed with water initiate fruit body formation for most species of morels 67 In 2021 it was announced that indoor cultivation of black morels had been successfully achieved after decades of research and experimentation with methods by The Danish Morel Project The project has been able to cultivate 20 lbs of morels per square yard or around 10 kg per square metre with cost estimates expected to be similar to producing white button mushrooms Agaricus bisporus Previous attempts at cultivation had managed to produce sclerotia but encountered issues in getting them to reliably fruit One of the breakthroughs with this project was growing them in a climate controlled environment in conjunction with grass which is involved in stimulating fruiting in the morel mycelium Cultivation in this manner has been noted to produce superior morels for culinary uses since they can be assured to be insect slug and dirt free and therefore do not need to be washed and cleaned like foraged morels Since washing morels can negatively impact the texture reliable cultivation may result in more versatility with this ingredient in the kitchen as well as making the delicacy more affordable and accessible 70 71 Toxicity editMorchella species are thought to contain small amounts of hydrazine 72 toxins or an unknown toxin that is destroyed through cooking the presence of hydrazine is controversial since there are no primary references of hydrazine having been detected in the species because of this morels should never be eaten raw 73 It has been reported that even cooked morels can sometimes cause symptoms of upset stomach when consumed with alcohol 74 When eating this fungus for the first time it is wise to consume a small amount to minimize any allergic reaction As with all fungi morels for consumption must be clean and free of decay Morels growing in old apple orchards previously treated with the now banned insecticide lead arsenate may accumulate levels of toxic lead and arsenic that are unsuitable for human consumption 75 Uses editMorels almost universally associated with spring can be found in many habitats Morel may be more likely to fruit during a period of increasing heat following a chilly period a preference which is credited for their abundance in areas with cold winters 76 Black morels Morchella elata are often found on land that has been disturbed by logging burning 76 Nutrition edit Morel mushrooms rawNutritional value per 100 g 3 5 oz Energy129 kJ 31 kcal Carbohydrates5 1 gSugars0 6 gDietary fiber2 8 gFat0 57 gProtein3 12 gVitaminsQuantity DV Thiamine B1 6 0 069 mgRiboflavin B2 17 0 205 mgNiacin B3 15 2 252 mgPantothenic acid B5 9 0 44 mgVitamin B610 0 136 mgFolate B9 2 9 mgVitamin D34 5 1 mgMineralsQuantity DV Calcium4 43 mgIron94 12 18 mgMagnesium5 19 mgManganese28 0 587 mgPhosphorus28 194 mgPotassium14 411 mgZinc21 2 03 mgOther constituentsQuantityWater90 gLink to USDA Database entryUnits mg micrograms mg milligrams IU International units Percentages are roughly approximated using US recommendations for adults Source USDA FoodData CentralRaw morel mushrooms are 90 water 5 carbohydrates 3 protein and 1 fat A 100 gram reference amount supplies 31 calories and is a rich source of iron 94 of the Daily Value DV manganese phosphorus zinc and vitamin D 34 DV if having been exposed to sunlight or artificial ultraviolet light Raw morels contain moderate levels of several B vitamins table Gastronomical value and culinary uses edit They have been called prized delicacies they are so esteemed in Europe that people used to set fire to their own forests in hopes of eliciting a bountiful morel crop the next spring 76 Morels are a feature of many cuisines including Provencal 77 Their flavor is prized by chefs worldwide with recipes and preparation methods designed to highlight and preserve it 78 As with most edible fungi they are best when collected or bought fresh They are sometimes added to meat and poultry dishes and soups and can be used as pasta fillings 79 As morels are known to contain thermolabile toxins they must always be cooked before eating Morels can be preserved in several ways They can be flash frozen by simply running under cold water or putting them in a bucket to soak for a few minutes then spread on a baking tray and placed into a freezer After freezing they keep very well with the frozen glaze for a long time in airtight containers However when thawed they can sometimes turn slightly mushy so they are best frozen after steaming or frying Due to their natural porosity morels may contain trace amounts of soil which cannot be easily washed out Any visible soil should be removed with a brush after cutting the body in half lengthwise if needed Mushroom hunters sometimes recommend soaking morels in a bowl of salt water briefly prior to cooking although many chefs would disagree 80 Drying is a popular and effective method for long term storage and morels are widely available commercially in this form Any insect larvae which might be present in the fruit bodies usually drop out during the drying process 81 Dried morels can then be reconstituted by soaking for 10 20 minutes in warm water or milk and the soaking liquid can be used as stock 82 The flavor of morels is not just appreciated by humans in Yellowstone National Park black morels are also known to be consumed by grizzly bears Ursus arctos horribilis 83 In popular culture editMorel hunting is a common springtime activity Mushroom collectors may carry a mesh collecting bag so the spores can scatter as one carries the harvest 80 Every spring hundreds of morel enthusiasts gather in Boyne City Michigan for the National Morel Mushroom Festival a century old event 84 As one observer stated if there is a modern North American reenactment of Geoffrey Chaucer s Canterbury Tales this is it 7 Other festivals and hunting competitions in North America include the Illinois State Morel Mushroom Hunting Championship the Ottawa Midwest Morel Fest and the Mesick Michigan Mushroom Festival 85 In the survival horror video game Cataclysm Dark Days Ahead morel mushrooms are featured as one of the many edible items that the survivors can gather in the forests They typically can be picked from underbrushes during summer and autumn and can be boiled fried or preserved via drying In the farming sim video game Stardew Valley morels are a consumable item that can be found in the Secret Woods or Forest Farm during spring Its player chooses to allow mushrooms to grow in the Farm Cave morels have an 8 1 chance of spawning each day regardless of season The item s description reads that it is sought after for its unique and nutty flavor Vernacular names edit Morchella species have been called by many local names some of the more colorful include dryland fish because when sliced lengthwise then breaded and fried their outline resembles the shape of a fish 86 hickory chickens as they are known in many parts of Kentucky and merkels or miracles based on folklore of how a mountain family was saved from starvation by eating morels In parts of West Virginia they are known as molly moochers muggins or muggles Due to the partial structural and textural similarity to some species of Porifera sponges other common names for any true morel are sponge mushroom and waffle mushroom In the Appalachian woodlands morels have also been called haystacks or snakeheads 80 The Finnish vernacular name huhtasieni refers to huhta area cleared for agriculture by the slash and burn method 87 The scientific name of the genus Morchella itself is thought to have derived from morchel 88 an old German word close to Mohre carrot or beet due to similarity in shape nbsp Yellow morels in France nbsp Black morels in British Columbia Canada nbsp Morel in a house garden in Ben Shemen IsraelSee also edit nbsp Fungi portalReferences edit Morchella Dill ex Pers Species Fungorum CAB International Retrieved 2014 12 11 Michelot D Toth B 1991 Poisoning by Gyromitra esculenta a review Journal of Applied Toxicology 11 4 235 243 doi 10 1002 jat 2550110403 PMID 1939997 S2CID 7994829 Bresinsky A Besl H 1990 A colour atlas of poisonous fungi Wolfe Publishing Ltd London Karlson Stiber C Persson H 2003 Cytotoxic fungi an overview Toxicon 42 4 339 349 doi 10 1016 S0041 0101 03 00238 1 PMID 14505933 Lampe KF 1979 Toxic fungi Annual Review of Pharmacology and Toxicology 19 1 85 104 doi 10 1146 annurev pa 19 040179 000505 PMID 378111 Kuo M 2007 100 Edible Mushrooms Ann Arbor Michigan The University of Michigan Press pp 55 58 ISBN 978 0 472 03126 9 a b Kuo M 2005 Morels Ann Arbor Michigan University of Michigan Press p 173 ISBN 978 0 472 03036 1 Kimbrough JW 1970 Current trends in the classification of discomycetes Botanical Review 36 2 91 161 doi 10 1007 bf02858958 S2CID 11463562 Korf RP 1972 Synoptic key to the genera of the Pezizales Mycologia 64 5 937 994 doi 10 2307 3758070 JSTOR 3758070 Masaphy S Zabari L Goldberg D Jander Shagug G Spring 2010 The complexity of Morchella systematics A case of the yellow morel from Israel PDF Fungi Magazine 3 2 14 18 a b c Loizides M 2017 Morels the story so far Field Mycology 18 2 42 53 doi 10 1016 j fldmyc 2017 04 004 Overholts LO 1934 The morels of Pennsylvania Proceedings of the Pennsylvania Academy of Science 8 108 114 Seaver FJ 1942 The North American cup fungi Operculates Hafner Publishing C New 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arbutiphila and M disparilis spp nov Mycological Progress 15 4 39 doi 10 1007 s11557 016 1180 1 S2CID 15163613 Taskin H Dogan H Buyukalaca S Clowez P Moreau P A O Donnell K 2016 Four new morel Morchella species in the elata subclade M sect Distantes from Turkey Mycotaxon 131 2 467 482 doi 10 5248 131 467 Clowez P Marcos Martinez J Sanjaume R Marques G Bellanger J M Moreau P A 2020 A survey of half free morels in Spain reveals a new species Morchella iberica sp nov Ascomycota Pezizales Ascomycete org 12 1 11 18 doi 10 25664 art 0291 Yoon CS Gessner RV Romano MA 1990 Population genetics and systematics of the Morchella esculenta complex Mycologia 82 2 227 235 doi 10 2307 3759851 JSTOR 3759851 Bunyard B A Nicholson M S Royse D J 1994 A systematic assessment of Morchella using RFLP analysis of the 28S ribosomal gene Mycologia 86 6 762 72 doi 10 2307 3760589 JSTOR 3760589 Bunyard BA Nicholson MS Royse DJ 1995 Phylogenetic resolution of Morchella Verpa andDisciotis Pezizales Morchellaceae based on restriction enzyme analysis of the 28S ribosomal RNA gene Experimental Mycology 19 3 223 233 doi 10 1006 emyc 1995 1027 PMID 7553270 Kuo M March 2006 Morel Data Collection Project Preliminary results Retrieved 2009 05 26 a b c d e f O Donnell K Rooney AP Mills GL Kuo M Weber NS Rehner SA 2011 Phylogeny and historical biogeography of true morels Morchella reveals an early Cretaceous origin and high continental endemism and provincialism in the Holarctic Fungal Genetics and Biology 48 3 252 265 doi 10 1016 j fgb 2010 09 006 PMID 20888422 a b c Taskin H Buyukalaca S Hansen K O Donnell K 2012 Multilocus phylogenetic analysis of true morels Morchella reveals high levels of endemics in Turkey relative to other regions of Europe Mycologia 104 2 446 461 doi 10 3852 11 180 PMID 22123659 S2CID 207685509 a b c Stefani FO Sokolski S Wurtz TL Piche Y Hamelin RC Fortin JA Berube JA 2010 Morchella tomentosa a unique belowground structure and a new clade of morels PDF Mycologia 102 5 1082 1088 doi 10 3852 09 294 PMID 20943507 S2CID 6103729 Archived from the original PDF on 2011 07 20 Retrieved 2011 03 15 a b McFarlane EM Pilz D Weber NS 2005 High elevation gray morels and other Morchella species harvested as non timber forest products in Idaho and Montana PDF Mycologist 19 2 62 68 CiteSeerX 10 1 1 542 4063 doi 10 1017 S0269915X0500203X Harald Kellner Carsten Renker amp Francois Buscot 2005 Species diversity within the Morchella esculenta group Ascomycota Morchellaceae in Germany and France PDF Organisms Diversity amp Evolution 5 2 101 107 doi 10 1016 j ode 2004 07 001 Archived from the original PDF on 2011 07 19 Retrieved 2011 03 15 a b Hatira Taskin Saadet Buyukalacaa Hasan Huseyin Doganb Stephen A Rehnerc Kerry O Donnell Aug 2010 A multigene molecular phylogenetic assessment of true morels Morchella in Turkey Fungal Genetics and Biology 47 8 672 682 doi 10 1016 j fgb 2010 05 004 PMID 20580850 a b Masaphy S Zabari L Goldberg D 2009 New long season ecotype of Morchella rufobrunnea from northern Israel PDF Micologia Aplicada International 21 2 45 55 ISSN 1534 2581 a b c Du XH Zhao Q O Donnell K Rooney AP Yang ZL 2012 Multigene molecular phylogenetics reveals true morels Morchella are especially species rich in China Fungal Genetics and Biology 49 6 455 469 doi 10 1016 j fgb 2012 03 006 PMID 22503770 Pildain MB Visnovsky SB Barroetavena C 2014 Phylogenetic diversity of true morels Morchella the main edible non timber product from native Patagonian forests of Argentina Fungal Biology 118 9 10 755 763 doi 10 1016 j funbio 2014 03 008 hdl 11336 38473 PMID 25209634 Kanwal HK Acharya K Ramesh G Reddy MS 2010 Molecular characterization of Morchella species from the Western Himalayan region of India Current Microbiology 62 4 1245 1252 doi 10 1007 s00284 010 9849 1 PMID 21188589 S2CID 1394087 a b c d Loizides M Gonou Z Fransuas G Drakopoulos P Sammut C Martinis A Bellanger JM 2021 Extended phylogeography of the ancestral Morchella anatolica supports preglacial presence in Europe and Mediterranean origin of morels Mycologia 113 3 559 573 doi 10 1080 00275514 2020 1869489 PMID 33734016 S2CID 232298567 Buscot F Roux J 1987 Association between living roots and ascocarps of Morchella rotunda Transactions of the British Mycological Society 89 2 249 252 doi 10 1016 s0007 1536 87 80162 6 Buscot F 1992 Mycorrhizal succession and morel biology In Read DJ Lewis DH Fitter AH Alexander IJ eds Mycorrhizas in Ecosystems Wallingford United Kingdom CAB International pp 220 224 ISBN 978 0 85198 786 6 a b Dahlstrom JL Smith JE Weber NS 2000 Mycorrhiza like interaction by Morchella with species of the Pinaceae in pure culture synthesis Mycorrhiza 9 5 279 285 doi 10 1007 PL00009992 S2CID 36119264 Tedersoo L May TW Smith ME 2010 Ectomycorrhizal lifestyle in fungi global diversity distribution and evolution of phylogenetic lineages Mycorrhiza 20 4 217 263 doi 10 1007 s00572 009 0274 x PMID 20191371 S2CID 3351967 Lincoff GH 1981 The Audubon Society Field Guide to North American Mushrooms p 326 Du XH Zhao Q Yang ZL 2015 A review on research advances issues and perspectives of morels Mycology 6 2 78 85 doi 10 1080 21501203 2015 1016561 PMC 6106076 PMID 30151316 Taskin H Dogan HH Buyukalaca S 2015 Morchella galilaea an autumn species from Turkey Mycotaxon 130 215 221 doi 10 5248 130 215 Loizides M 2011 Morchella rufobrunnea h morxella ths polhs Morchella rufobrunnea the urban morel Mykhtologos Mycologist in Greek 5 10 13 a b c Pilz D R McLain S Alexander L Villarreal Ruiz S Berch T L Wurtz C G Parks E McFarlane B Baker R Molina J E Smith March 2007 Ecology and management of morels harvested from the forests of western North America General Technical Report PNW GTR 710 Portland OR U S Forest Service Pacific Northwest Research Station Mihail JD Bruhn JN Bonello P 2007 Spatial and temporal patterns of morel fruiting Mycological Research 111 3 339 346 doi 10 1016 j mycres 2007 01 007 PMID 17363234 a b Greene DF Hesketh M Pounden E 2010 Emergence of morel Morchella and pixie cup Geopyxis carbonaria ascocarps in response to the intensity of forest floor combustion during a wildfire Mycologia 102 4 766 773 doi 10 3852 08 096 PMID 20648745 S2CID 28400225 Wurtz TL Wiita AL Weber NS Pilz D 2005 Harvesting morels after wildfire in Alaska Report Research Note RN PNW 546 Portland Oregon U S Forest Service Pacific Northwest Research Station Archived from the original on 2011 09 28 Retrieved 2009 05 26 a b Archived at Ghostarchive and the Wayback Machine YouTube YouTube Growing Morels 2019 11 08 Retrieved 2019 11 11 Du XH Zhao Q Yang ZL Hansen K Taskin H Buyukalaca S Dewsbury D Moncalvo JM Douhan GW Robert VA Crous PW Rehner SA Rooney AP Sink S O Donnell K 2012 How well do ITS rDNA sequences differentiate species of true morels Morchella Mycologia 104 6 1351 1368 doi 10 3852 12 056 PMID 22802394 S2CID 6740906 Repin C 1901 Sur la culture de la Morille Revue generale des sciences pures et appliquees 12 595 96 Retrieved 2010 03 21 The Morel Project The Danish Morel Project 2021 11 16 Retrieved 2022 07 12 Gamillo Elizabeth 2022 05 02 Danish Biologists Cultivate Morel Mushrooms Year Round With New Indoor Technique Smithsonian Magazine Retrieved 2022 07 12 Stamets P 2005 Mycelium Running Berkeley Ten Speed Press p 344 Hall IR Buchanan PK 2003 Edible and Poisonous Mushrooms of the World Timber Press ISBN 978 0 88192 586 9 Groves JW 1964 Poisoning by morels When taken with alcohol Mycologia 56 5 779 780 doi 10 2307 3756634 JSTOR 3756634 Shavit E Shavit E Spring 2010 Lead and arsenic in Morchella esculenta fruitbodies collected in lead arsenate contaminated apple orchards in the northeastern United States A preliminary study PDF Fungi Magazine 3 2 11 18 a b c Arora David 1986 Mushrooms demystified a comprehensive guide to the fleshy fungi 2nd ed Berkeley Ten Speed Press pp 785 793 ISBN 0 89815 170 8 OCLC 13702933 Olney R 1995 A Provencal Table London Pavilion pp 31 32 ISBN 978 1 85793 632 2 Acton J Sandler N 2007 Mushroom Watermill Books p 160 ISBN 978 1856267397 Wright J 2007 Mushrooms River Cottage Cookbook 1 Bloomsbury p 256 ISBN 9780747589327 a b c Excerpts from the book Animal Vegetable Miracle by Barbara Kingsolver Animal Vegetable Miracle Harper Perennial pp 77 80 Wild About Mushrooms Morels Mssf org Retrieved on 2012 04 17 Carluccio A 2003 The Complete Mushroom Book London UK Quadrille pp 37 38 96 97 ISBN 978 1 84400 040 1 Mattson DJ Podruzny SR Haroldson MA 2002 Consumption of fungal sporocarps by Yellowstone grizzly bears Ursus 13 95 103 JSTOR 3873191 National Morel Mushroom Festival bcmorelfestival com Retrieved 2018 08 10 Annual Mesick Mushroom Festival Lincoff G 2011 The Complete Mushroom Hunter An Illustrated Guide to Finding Harvesting and Enjoying Wild Mushrooms Quarry Books p 57 ISBN 978 1 61058 099 1 Cunningham AB Yang X 2012 Mushrooms in Forests and Woodlands Resource Management Values and Local Livelihoods Routledge p 180 ISBN 978 1 136 53817 9 Burnett GT 1835 Outlines of Botany Including a General History of the Vegetable Kingdom in which Plants are Arranged According to the System of Natural Affinities John Churchill p 241 External links edit nbsp Wikimedia Commons has media related to Morchella 2000 Pezizales on Ascomycete org MushroomExpert com s Morel section A Beginner s Guide to Hunting Morel Mushrooms from Field and Stream Colorado Morels Archived 2020 11 11 at the Wayback Machine Retrieved from https en wikipedia org w index php title Morchella amp oldid 1191997824, wikipedia, wiki, book, books, library,

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