''Morchella'', the true morels, is a

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

of edible sac fungi closely related to anatomically simpler cup fungi in the order

Pezizales The Pezizales are an order of the subphylum Pezizomycotina within the phylum Ascomycota. The order contains 16 families, 199 genera, and 1683 species. It contains a number of species of economic importance, such as morels, the black and whit ...

( division

Ascomycota Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...

). These distinctive

fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...

have a

honeycomb A honeycomb is a mass of Triangular prismatic honeycomb#Hexagonal prismatic honeycomb, hexagonal prismatic cells built from beeswax by honey bees in their beehive, nests to contain their brood (eggs, larvae, and pupae) and stores of honey and pol ...

appearance due to the network of ridges with pits composing their caps. Morels are prized by

gourmet Gourmet (, ) is a cultural idea associated with the culinary arts of fine food and drink, or haute cuisine, which is characterized by their high level of refined and elaborate food preparation techniques and displays of balanced meals that have ...

cooks, particularly in Catalan and

French cuisine French cuisine is the cooking traditions and practices of France. In the 14th century, Guillaume Tirel, a Court (royal), court chef known as "Taillevent", wrote ''Le Viandier'', one of the earliest recipe collections of medieval France. In ...

, but can be toxic if consumed raw or undercooked. Due to difficulties in cultivation, commercial harvesting of wild morels has become a multimillion-dollar industry in the temperate

Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...

, in particular

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

Turkey Turkey, officially the Republic of Türkiye, is a country mainly located in Anatolia in West Asia, with a relatively small part called East Thrace in Southeast Europe. It borders the Black Sea to the north; Georgia (country), Georgia, Armen ...

China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...

, the

Himalayas The Himalayas, or Himalaya ( ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the Earth's highest peaks, including the highest, Mount Everest. More than list of h ...

India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...

, and

Pakistan Pakistan, officially the Islamic Republic of Pakistan, is a country in South Asia. It is the List of countries and dependencies by population, fifth-most populous country, with a population of over 241.5 million, having the Islam by country# ...

where these highly prized fungi are found in abundance. 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 Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

suggest there might be over seventy species of ''Morchella'' worldwide, most of them exhibiting high continental

endemism Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

and provincialism. The genus is currently the focus of extensive

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

, biogeographical, taxonomical and nomenclatural studies, and several new species have been described from Australia, Canada, Cyprus, Israel, Spain, and Turkey.

Description

Morels resemble a

due to the network of ridges with pits composing their caps. Morels have a convoluted head/cap, and are varied in shape and habitat. File:Morchella americana on a mossy riverbank in MI.jpg File:Morchella_americana_closer.JPG File:Morchella esculenta 5.jpg File:Morchella esculenta 6.jpg File:Old holiday shot of Morchella esculenta (GB= Morel mushroom, D= Speise-Morchel, NL= Gewone morielje) - panoramio.jpg File:Morchella esculenta Illinois.jpg

Similar species

When foraging for morels, one must be absolutely sure of identification. There are many look-alikes often referred to as " false morels", most notably '' Gyromitra''. These also include members of the most closely related genus, '' Verpa'', which are highly poisonous. Other mushrooms can also be mistaken for morels, including some species of stinkhorns, or

Phallaceae Phallaceae is a family of fungi, commonly known as stinkhorns, within the order Phallales. Stinkhorns have a worldwide distribution, but are especially prevalent in tropical regions. They are known for their foul-smelling, sticky spore masses ...

, which have a similarly shaped cap but a distinctive foul odor. It is important to take care when harvesting and identifying mushrooms, particularly morels. 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. * The caps of morels come in a range of colors including white, grey, black, brown, and yellow. ''Gyromitra esculenta'' has a cap that is usually reddish-brown in colour, but sometimes also chestnut, purplish-brown, or dark brown. * True morels are always hollow when sliced lengthwise, whereas ''Gyromitra'' species are typically chambered in longitudinal sections, while '' Verpa'' species contain a cottony substance inside their stem. The easiest way to distinguish ''Verpa'' species from ''Morchella'' species is to slice them longitudinally ,8 * 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.

Taxonomy

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. 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,Weber, N.S. (1988). In A Morel Hunter's Companion, pp. 111-167. Two Peninsula Press: Lansing. while others recognised as many as 34. 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 A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

material. In 2012, the simultaneous description of several new taxa from Europe by Clowez and North America by Kuo and colleagues resulted in several

synonyms A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...

further complicating matters, until a transatlantic study by Richard and colleagues resolved many of these issues in 2014. The genus is currently undergoing extensive re-evaluation with regard to the taxonomic status of several species.

Early taxonomic history

''Morchella'' Dill. ex Pers. : Fr. was typified by

Christiaan Hendrik Persoon Christiaan Hendrik Persoon (31 December 1761 – 16 November 1836) was a Cape Colony mycologist who is recognized as one of the founders of mycology, mycological Taxonomy (biology), taxonomy. Early life Persoon was born in Cape Colony at ...

in 1794, with '' Morchella esculenta'' designated as the

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

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 and 1897 respectively, published several species and varieties, accompanied by meticulously illustrated iconographic plates. The seminal taxon '' Morchella elata'', whose true identity still remains unresolved, was described by

Elias Fries Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist and botanist. He is sometimes called the "Linnaeus of Mycology". In his works he described and assigned botanical names to hundreds of fungus and lichen sp ...

in 1822, from a fir forest in

Sweden Sweden, formally the Kingdom of Sweden, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. It borders Norway to the west and north, and Finland to the east. At , Sweden is the largest Nordic count ...

. 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, '' Morchella vulgaris'', which was recombined by Samuel Gray as a distinct species in 1821 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. '' 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. '' Morchella eximia'', a globally-occurring fire-associated species was also described by Boudier in 1910. The old, widely applied name '' Morchella conica'', featuring in many field guides and literature across several countries, has been shown by Richard and colleagues to be illegitimate.

Classification

About 80 species of ''Morchella'' were described until the turn of the 21st century (per the ''

Index Fungorum ''Index Fungorum'' is an international project to index all formal names (scientific names) in the fungus kingdom. As of 2015, the project is based at the Royal Botanic Gardens, Kew, one of three partners along with Landcare Research and th ...

''), a number of which were later shown to be illegitimate or synonyms. 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. In 2010 Işiloğlu and colleagues described '' Morchella anatolica'', a basal species from

later shown to be sister to '' Morchella rufobrunnea''. A study by Clowez described over 20 new species in 2012, while later in the same year, another study by Kuo and colleagues described 19 species from

. However, several of these newly proposed names later turned out to be synonyms. 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. Also in 2014, Elliott and colleagues described '' Morchella australiana'' from

sclerophyll Sclerophyll is a type of vegetation that is adapted to long periods of dryness and heat. The plants feature hard leaves, short Internode (botany), internodes (the distance between leaves along the stem) and leaf orientation which is parallel or ...

forests in

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

, while Clowez and colleagues described '' Morchella fluvialis'' from riparian forests in

Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...

. 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. Later in the same year, Clowez and colleagues described '' Morchella palazonii'' from Spain, while Voitk and colleagues described '' Morchella laurentiana'' from

Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...

and '' Morchella eohespera'', a cosmopolitan species present in several continents. In an extensive phylogenetic and morphological study from

Cyprus Cyprus (), officially the Republic of Cyprus, is an island country in the eastern Mediterranean Sea. Situated in West Asia, its cultural identity and geopolitical orientation are overwhelmingly Southeast European. Cyprus is the List of isl ...

in 2016, Loizides and colleagues added two more

Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...

species, '' Morchella arbutiphila'' and '' Morchella disparilis'', and resurrected '' Morchella dunensis'' as an autonomous species. 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''.

Phylogeny

Early

analyses supported the hypothesis that the genus comprises only a few species with considerable

phenotypic In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...

variation. Subsequent multigenic DNA studies, however, have revealed more than a dozen genealogically distinct species in

and at least as many in

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

. DNA studies revealed three discrete

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

s, or genetic groups, consisting of the "white morels" ('' Morchella rufobrunnea'' and '' M. anatolica''), the "yellow morels" ('' M. esculenta'' and others), and the "black morels" ('' M. elata'' and others). 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. Within the yellow and black clades, there are dozens of distinct species, many

endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

to individual continents or regions. This species-rich view is supported by studies in

Western Europe Western Europe is the western region of Europe. The region's extent varies depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the Western half of the ancient Mediterranean ...

, Turkey,

Israel Israel, officially the State of Israel, is a country in West Asia. It Borders of Israel, shares borders with Lebanon to the north, Syria to the north-east, Jordan to the east, Egypt to the south-west, and the Mediterranean Sea to the west. Isr ...

Patagonia Patagonia () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...

, and the

. Early

ancestral reconstruction Ancestral reconstruction (also known as ''Character Mapping'' or ''Character Optimization'') is the extrapolation back in time from measured characteristics of individuals, populations, or species to their Common descent, common ancestors. It is an ...

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). This date was later revised by Du and collaborators, placing the divergence of the genus in the late Jurassic, approximately 154 Mya. 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 ''M. rufobrunnea''. 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. 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.

Distribution and habitat

Morels can be found in the temperate

, in particular North America, Turkey, China, the Himalayas,

, and

Yellow morels ('' Morchella esculenta'' and related species) are more commonly found under

deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...

trees rather than

conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...

s, while black morels ('' M. elata'' and related species) are mostly found in coniferous forests,

disturbed ground In ecology, a disturbance is a change in environmental conditions that causes a pronounced change in an ecosystem. Disturbances often act quickly and with great effect, to alter the physical structure or arrangement of biotic and abiotic element ...

and recently burned areas. '' Morchella galilaea'', and occasionally ''M. rufobrunnea'', appear to fruit in the autumn or winter months rather than spring, which is the typical fruiting season for morels. In the American

Pacific Northwest The Pacific Northwest (PNW; ) is a geographic region in Western North America bounded by its coastal waters of the Pacific Ocean to the west and, loosely, by the Rocky Mountains to the east. Though no official boundary exists, the most common ...

, they can be found from April to August. Efforts to cultivate morels at a large scale have rarely been successful and the commercial morel industry relies on the harvest of wild mushrooms.

Transcontinental species

Although many species within ''Morchella'' exhibit continental

and provincialism, 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''. 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, 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. Long-distance spore dispersal has also been suggested as a possible dispersal mechanism for some species, especially those belonging to fire-adapted lineages. 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 The Quaternary glaciation, also known as the Pleistocene glaciation, is an alternating series of glacial period, glacial and interglacial, interglacial periods during the Quaternary period that began 2.58 Year#SI prefix multipliers, Ma (million ...

Ecology

The ecology of ''Morchella'' species is not well understood. Many species appear to form

symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...

or endophytic relationships with trees, while others appear to act as

saprotroph Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ...

s. 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

include '' Abies'' (fir), ''

Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as c ...

'' (pine), ''

Populus ''Populus'' is a genus of 25–30 species of deciduous flowering plants in the family Salicaceae, native to most of the Northern Hemisphere. English names variously applied to different species include poplar (), aspen, and cottonwood. The we ...

'' (poplar), ''

Ulmus Elms are deciduous and semi-deciduous trees comprising the genus ''Ulmus'' in the family Ulmaceae. They are distributed over most of the Northern Hemisphere, inhabiting the temperate and tropical- montane regions of North America and Eurasia, ...

'' (elm), ''

Quercus An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...

'' (oak), '' Arbutus'' (strawberry trees), '' Castanea'' (chestnut), '' Alnus'' (alder), '' Olea'' (olive trees), ''

Malus ''Malus'' ( or ) is a genus of about 32–57 species of small deciduous trees or shrubs in the family Rosaceae, including the domesticated orchard apple, crab apples (sometimes known in North America as crabapples) and wild apples. The genus i ...

'' (apple trees), and ''

Fraxinus ''Fraxinus'' (), commonly called ash, is a genus of plants in the olive and lilac family, Oleaceae, and comprises 45–65 species of usually medium-to-large trees, most of which are deciduous trees, although some Subtropics, subtropical specie ...

'' (ash). In western North America morels are often found in coniferous forests, including species of ''

'' (pine), '' Abies'' (fir), '' Larix'' (larch), and '' Pseudotsuga'' (Douglas-fir), as well as in ''

'' (cottonwood)

riparian A riparian zone or riparian area is the interface between land and a river or stream. In some regions, the terms riparian woodland, riparian forest, riparian buffer zone, riparian corridor, and riparian strip are used to characterize a ripar ...

forests. Deciduous trees commonly associated with morels in the northern hemisphere include ''

'' (ash), ''

Platanus ''Platanus'' ( ) is a genus consisting of a small number of tree species native to the Northern Hemisphere. They are the sole living members of the family Platanaceae. All mature members of ''Platanus'' are tall, reaching in height. The type ...

'' (sycamore), ''

Liriodendron ''Liriodendron'' () is a genus of two species of characteristically large trees, deciduous tree, deciduous over most of their populations, in the magnolia family (biology), family (Magnoliaceae). These trees are widely known by the common name ...

'' (tulip tree), dead and dying elms, cottonwoods, and old apple trees (remnants of

orchard An orchard is an intentional plantation of trees or shrubs that is maintained for food production. Orchards comprise fruit tree, fruit- or nut (fruit), nut-producing trees that are generally grown for commercial production. Orchards are also so ...

s). Due to their springtime

phenology Phenology is the study of periodic events in biological life cycles and how these are influenced by seasonal and interannual variations in climate, as well as habitat factors (such as elevation). Examples include the date of emergence of leav ...

(March–May), morels are hardly ever found in the vicinity of common poisonous mushrooms such as the

death cap ''Amanita phalloides'' ( ), commonly known as the death cap, is a deadly poisonous basidiomycete fungus and mushroom, one of many in the genus ''Amanita''. Originating in Europe but later introduced to other parts of the world since the late ...

(''Amanita phalloides''), the sulphur tuft (''Hypholoma fasciculare''), or the

fly agaric ''Amanita muscaria'', commonly known as the fly agaric or fly amanita, is a basidiomycete fungus of the genus ''Amanita''. It is a large white- gilled, white-spotted mushroom typically featuring a bright red cap covered with distinctive white ...

(''Amanita muscaria''). They can, however, occur alongside false morels ('' Gyromitra'' and '' Verpa'' species) and elfin saddles ('' Helvella'' species), which also appear in spring.

Association with wildfire

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 Fire is the rapid oxidation of a fuel in the exothermic chemical process of combustion, releasing heat, light, and various reaction Product (chemistry), products. Flames, the most visible portion of the fire, are produced in the combustion re ...

. Moderate-intensity fires are reported to produce higher abundances of morels than low- or high-intensity fires. This is caused by the soil becoming more alkaline as the result of

wood ash Wood ash is the powder (substance), powdery residue remaining after the combustion of wood, such as burning wood in a fireplace, bonfire, or an industrial power plant. It is largely composed of calcium compounds, along with other non-combustible ...

combining with water and being absorbed into the soil which triggers the morels to fruit.

Alkaline soil Alkali, or alkaline, soils are clay soils with high Soil pH, pH (greater than 8.5), a poor soil structure and a low infiltration capacity. Often they have a hard calcareous layer at 0.5 to 1 metre depth. Alkali soils owe their unfavorable ...

conditions which trigger fruiting have been observed and exploited with small-scale commercial cultivation of morels.Archived a
Ghostarchive
and th
Wayback Machine
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 A cash crop, also called profit crop, is an Agriculture, agricultural crop which is grown to sell for profit. It is typically purchased by parties separate from a farm. The term is used to differentiate a marketed crop from a staple crop ("subsi ...

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. 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. 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 ''Agaricus bisporus'', commonly known as the cultivated mushroom, is a basidiomycete mushroom native to grasslands in Eurasia and North America. It is cultivated in more than 70 countries and is one of the most commonly and widely consumed ...

''). 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.

Toxicity

The consumption of ''Morchella'' species can have adverse effects. In 2023, a Montana

sushi is a traditional Japanese dish made with , typically seasoned with sugar and salt, and combined with a variety of , such as seafood, vegetables, or meat: raw seafood is the most common, although some may be cooked. While sushi comes in n ...

restaurant serving them was linked to 51 people who experienced

gastrointestinal illness Gastrointestinal diseases (abbrev. GI diseases or GI illnesses) refer to diseases involving the gastrointestinal tract, namely the esophagus, stomach, small intestine, large intestine and rectum; and the accessory organs of digestion, the liver, ...

, with two reported deaths and three other hospitalizations. The consumption of raw morels in particular is advised against. An unknown toxin can be neutralized via cooking. Additionally, cooked morels can reportedly cause upset stomachs when consumed with alcohol. When eating this fungus for the first time, it is advised 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 An apple is a round, edible fruit produced by an apple tree (''Malus'' spp.). Fruit trees of the orchard or domestic apple (''Malus domestica''), the most widely grown in the genus, are agriculture, cultivated worldwide. The tree originated ...

orchards previously treated with the deprecated insecticide lead arsenate may accumulate levels of toxic

lead Lead () is a chemical element; it has Chemical symbol, symbol Pb (from Latin ) and atomic number 82. It is a Heavy metal (elements), heavy metal that is density, denser than most common materials. Lead is Mohs scale, soft and Ductility, malleabl ...

and

arsenic Arsenic is a chemical element; it has Symbol (chemistry), symbol As and atomic number 33. It is a metalloid and one of the pnictogens, and therefore shares many properties with its group 15 neighbors phosphorus and antimony. Arsenic is not ...

that are unsuitable for human consumption.

Uses

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

Nutrition

Raw morel mushrooms are 90% water, 5%

carbohydrate A carbohydrate () is a biomolecule composed of carbon (C), hydrogen (H), and oxygen (O) atoms. The typical hydrogen-to-oxygen atomic ratio is 2:1, analogous to that of water, and is represented by the empirical formula (where ''m'' and ''n'' ...

s, 3%

protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...

, and 1%

fat In nutrition science, nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such chemical compound, compounds, most commonly those that occur in living beings or in food. The term often refers specif ...

. A 100 gram reference amount supplies 31

calorie The calorie is a unit of energy that originated from the caloric theory of heat. The large calorie, food calorie, dietary calorie, kilocalorie, or kilogram calorie is defined as the amount of heat needed to raise the temperature of one liter o ...

s, and is a rich source of

iron Iron is a chemical element; it has symbol Fe () and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, forming much of Earth's o ...

(94% of the Daily Value, DV),

manganese Manganese is a chemical element; it has Symbol (chemistry), symbol Mn and atomic number 25. It is a hard, brittle, silvery metal, often found in minerals in combination with iron. Manganese was first isolated in the 1770s. It is a transition m ...

phosphorus Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...

zinc Zinc is a chemical element; it has symbol Zn and atomic number 30. It is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodic tabl ...

, and vitamin D (34% DV, if having been exposed to sunlight or artificial ultraviolet light). Raw morels contain moderate levels of several

B vitamins B vitamins are a class of water-soluble vitamins that play important roles in Cell (biology), cell metabolism and synthesis of red blood cells. They are a chemically diverse class of compounds. Dietary supplements containing all eight are referr ...

(table).

Gastronomical value and culinary uses

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!" Morels are a feature of many cuisines, including Provençal. Their flavor is prized by

chef A chef is a professional Cook (profession), cook and tradesperson who is proficient in all aspects of outline of food preparation, food preparation, often focusing on a particular cuisine. The word "chef" is derived from the term (), the di ...

s worldwide, with recipes and preparation methods designed to highlight and preserve it. 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 Pasta (, ; ) is a type of food typically made from an Leavening agent, unleavened dough of wheat flour mixed with water or Eggs as food, eggs, and formed into sheets or other shapes, then cooked by boiling or baking. Pasta was originally on ...

fillings. As morels are known to contain

thermolabile Thermolabile refers to a substance which is subject to decomposition or change in response to heat. This term is often used describe biochemical substances. For example, many bacterial exotoxins are thermolabile and can be easily inactivate ...

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 Steaming is a method of cooking using steam. This is often done with a food steamer, a kitchen appliance made specifically to cook food with steam, but food can also be steamed in a wok. In the American Southwest, steam pits used for cooking ha ...

frying Frying is the cooking of food in cooking oil, oil or another fat. Similar to sautéing, pan-fried foods are generally turned over once or twice during cooking to make sure that the food is evenly cooked, using tongs or a spatula, whilst sautéed ...

. 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. 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.Wild About Mushrooms: Morels
Mssf.org. Retrieved on 2012-04-17. 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. The flavor of morels is not just appreciated by humans; in

Yellowstone National Park Yellowstone National Park is a List of national parks of the United States, national park of the United States located in the northwest corner of Wyoming, with small portions extending into Montana and Idaho. It was established by the 42nd U ...

, black morels are also known to be consumed by

grizzly bear The grizzly bear (''Ursus arctos horribilis''), also known as the North American brown bear or simply grizzly, is a population or subspecies of the brown bear inhabiting North America. In addition to the mainland grizzly (''Ursus arctos horr ...

s (''Ursus arctos horribilis'').

In popular culture

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. Every spring, hundreds of morel enthusiasts gather in

Boyne City, Michigan Boyne City () is a city in Charlevoix County, Michigan, Charlevoix County in the U.S. state of Michigan. The population was 3,816 at the 2020 United States census, 2020 census, making it the largest city in Charlevoix County. Boyne City is locat ...

for the National Morel Mushroom Festival, a century-old event. As one observer stated, "if there is a modern, North American reenactment of

Geoffrey Chaucer Geoffrey Chaucer ( ; – 25 October 1400) was an English poet, author, and civil servant best known for ''The Canterbury Tales''. He has been called the "father of English literature", or, alternatively, the "father of English poetry". He w ...

's ''

Canterbury Tales ''The Canterbury Tales'' () is a collection of 24 stories written in Middle English by Geoffrey Chaucer between 1387 and 1400. The book presents the tales, which are mostly written in verse (poetry), verse, as part of a fictional storytellin ...

'' this is it." Other festivals and hunting competitions in

include the Illinois State Morel Mushroom Hunting Championship, the Ottawa Midwest Morel Fest and the Mesick Michigan Mushroom Festival. In Tyler Childers 2019 song " All Your'n", he mentions "Fried morels and fine hotels" in the first line of the 2nd verse. In the farming sim video game '' Stardew Valley'', morels are a consumable item that can be found in the Secret Woods during spring. If the player chooses to allow mushrooms to grow in the Farm Cave, morels have an 8.1% chance of spawning each day regardless of season.

Vernacular names

''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; ''hickory chickens'', as they are known in many parts of

Kentucky Kentucky (, ), officially the Commonwealth of Kentucky, is a landlocked U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders Illinois, Indiana, and Ohio to the north, West Virginia to the ...

; and ''merkels'' or ''miracles'', based on

folklore Folklore is the body of expressive culture shared by a particular group of people, culture or subculture. This includes oral traditions such as Narrative, tales, myths, legends, proverbs, Poetry, poems, jokes, and other oral traditions. This also ...

, of how a mountain family was saved from starvation by eating morels. In parts of

West Virginia West Virginia is a mountainous U.S. state, state in the Southern United States, Southern and Mid-Atlantic (United States), Mid-Atlantic regions of the United States.The United States Census Bureau, Census Bureau and the Association of American ...

, they are known as ''molly moochers'', ''muggins'', or ''muggles''. Due to the partial structural and textural similarity to some species of ''

Porifera Sponges or sea sponges are primarily marine invertebrates of the animal phylum Porifera (; meaning 'pore bearer'), a Basal (phylogenetics) , basal clade and a sister taxon of the Eumetazoa , diploblasts. They are sessility (motility) , sessile ...

'' (sponges), other common names for any true morel are ''sponge mushroom'' and ''

waffle A waffle is a dish made from leavened Batter (cooking), batter or dough that is cooked between two plates that are patterned to give a characteristic size, shape, and surface impression. There are many variations based on the type of waffle iron ...

mushroom''. In the Appalachian woodlands, morels have also been called ''haystacks'', or ''snakeheads''. The Finnish vernacular name ''huhtasieni'', refers to ''huhta'', area cleared for agriculture by the slash and burn method. The scientific name of the genus ''Morchella'' itself, is thought to have derived from ''morchel'', an old German word close to "Möhre", carrot or beet, due to similarity in shape. File:Morilles.jpg, left, Yellow morels in

France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...

File:Morchella elata group 38336.jpg, Black morels in

British Columbia British Columbia is the westernmost Provinces and territories of Canada, province of Canada. Situated in the Pacific Northwest between the Pacific Ocean and the Rocky Mountains, the province has a diverse geography, with rugged landscapes that ...

, Canada File:Morel in Ben Shemen Israel.jpg, Morel in a house garden in Ben Shemen, Israel

References

External links

2000 Pezizales
on Ascomycete.org

"A Beginner's Guide to Hunting Morel Mushrooms"
from ''Field and Stream''
Colorado Morels
{{Authority control Morel Pezizales genera