Brief Summary

The true morels (Morchella spp.) are among the edible fungi most prized by mushroom hunters.  As with other mushrooms, the familiar morel is merely the spore-producing "fruiting body" of a macrofungus which exists mostly undergound. A variety of morel species fruit briefly and sporadically each spring across the northern hemisphere and information on when and where to find them is often closely guarded by collectors. Much lore exists (some of it surely well founded) about the microhabitats and weather conditions associated with the appearance of morels. Morels grow throughout the northern hemisphere in regions with temperate or boreal forests, as well as in some Mediterranean and subtropical regions such as coastal California, the highlands of Central American, and the Middle East. Morels also occur in the southern hemisphere and although many of these are believed to be introduced, there are apparently endemic species as well in, for example, Australia and southern South America. Morels are harvested from the wild commercially in several parts of the world, including the United States, Turkey, China, and the Indian subcontinent, although some progress has been made toward commercial cultivation. (Pilz et al. 2007 and references therein). Pilz et al. (2007) provide an overview of the biology and ecology of morels.

In recent years, a number of researchers have used molecular genetic approaches to help resolve questions about species boundaries and species diversity within the genus Morchella (e.g., O'Donnell et al. 2011; Du et al. 2012).  Du et al. (2012) recognized more than five dozen putative Morchella species. Building on this work, Kuo et al. (2012) formally described a number of new phylogenetic species from the U.S. and Canada and reviewed their current understanding of the taxonomy and nomenclature of Morchella in the U.S. and Canada (Kuo et al. 2012 includes a dichotomous identification key). Much taxonomic work remains to be done on this challenging group, but it appears that the old idea that Morchella includes just a handful of very widely distributed species is unlikely to persist in the face of much new data indicating high levels of genetic diversity within geographic regions.

Although they are widely considered to be choice edibles and large numbers are eaten each year without ill effect, morels reportedly can be toxic, especially when poorly cooked, producing both gastrointestinal and neurological symptoms. Researchers investigating reports of Morchella toxicity concluded that these cases were not simply the result of confusion with the superficially similar and poisonous False Morel (Gyromitra esculenta) or other causes, but noted the need for further study to confirm and better understand the phenomenon of morel poisoning. (Saviuc et al 2010)

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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
Specimen Records:468
Specimens with Sequences:461
Specimens with Barcodes:457
Species With Barcodes:75
Public Records:446
Public Species:73
Public BINs:0
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Barcode data

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"Morel" redirects here. For other uses, see Morel (disambiguation).
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
Thiamine (B1)
0.069 mg
Riboflavin (B2)
0.205 mg
Niacin (B3)
2.252 mg
0.44 mg
Vitamin B6
0.136 mg
Folate (B9)
9 μg
Vitamin D
5.1 μg
Trace metals
43 mg
12.18 mg
19 mg
0.587 mg
194 mg
411 mg
2.03 mg

Percentages are roughly approximated using US recommendations for adults.
Source: USDA Nutrient Database

Morchella, the true morels, is a genus of edible mushrooms closely related to anatomically simpler cup fungi. These distinctive mushrooms appear honeycomb-like in that the upper portion is composed of a network of ridges with pits between them. The ascocarps are prized by gourmet cooks, particularly for French cuisine. Commercial value aside, morels are hunted by thousands of people every year simply for their taste and the joy of the hunt.

Morels 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; and merkels or miracles, based on a story of how a mountain family was saved from starvation by eating morels. In parts of West Virginia, they are known as molly moochers. Due to the partial structural and textural similarity to some species of the Porifera sponges, a common name for any true morel is sponge mushroom. Genus Morchella is derived from morchel, an old German word for mushroom, while morel itself is derived from the Latin maurus meaning brown.



Black morels in British Columbia, Canada

The fruit bodies of the Morchella are highly polymorphic in appearance, exhibiting variations in shape, color and size; this has contributed to uncertainties regarding taxonomy.[2] Discriminating between the various species is complicated by uncertainty regarding which species are truly biologically distinct. Some authors suggest that the genus only contains as few as 3 to 6 species,[3][4] while others place up to 50 species in the genus.[5][6] Mushroom hunters refer to them by their color (e.g., gray, yellow, black) as the species are very similar in appearance and vary considerably within species and age of individual. The best known morels are the "yellow morel" or "common morel" (M. esculenta); the "white morel" (M. deliciosa); and the "black morel" (M. elata). Other species of true morels include M. conica, M. vulgaris, and the half-free morel (M. semilibera).


Early phylogenetic analyses supported the hypothesis that the genus comprises only a few species with considerable phenotypic variation.[7][8] More recent DNA work has suggested more than a dozen distinct groups of morels in North America.[9] An extensive DNA study showed three discrete clades, or genetic groups, consisting of Morchella rufobrunnea, the yellow morels (M. esculenta and others), and the black morels (M. elata and others). Within the yellow and black clades, there are dozens of individual species, most endemic to individual continents or regions.[10] This species-rich view is supported by studies in Western Europe,[11] Turkey,[12] Israel,[13] and the Himalayas.[14]

Morchella tomentosa, a fire-associated species described from western North America, commonly known as the "gray morel", may also deserve its own clade based on DNA evidence. M. tomentosa is easily identified by its post-fire occurrence, fine hairs on the surface of young fruiting bodies, and unique sclerotia-like underground parts.[15][16][17]


A 2012 study described 19 phylogenetic species that occur in North America,[18] while molecular phylogenetics suggest that there are more than 60 species of Morchella worldwide.[19] Over 20 new species were added to the genus in 2012 by Clowez and colleagues.[20] A further revision of the taxonomy of the genus was provided by Richard et al. in 2014.[21]

Section Rufobrunnea[edit]

Section Morchella[edit]

Section Distantes[edit]

Uncertain classification[edit]

Habitat and ecology[edit]

Longneck morel in Indiana, USA
Yellow morels in West Virginia, USA

Black morels are known to be consumed by grizzly bears (species Ursus arctos horribilis) in Yellowstone National Park.[23]

Habitats favorable to fruition[edit]

Morchella species appear to have either symbiotic mycorrhizal relationships or act as saprotrophs.[15][24] Yellow morels (Morchella esculenta) are more commonly found under deciduous trees rather than conifers, and black morels (Morchella elata) can be found in deciduous forests, oak and poplar.[25] Deciduous trees commonly associated with morels in the northern hemisphere include ash, sycamore, tulip tree, dead and dying elms, cottonwoods and old apple trees (remnants of orchards). The fruiting of yellow morels in Missouri, USA, was found to correlate with warm weather, precipitation, and tree species, and most usually in the springtime. In the UK, they appear during May and June. Morels are rarely found in the vicinity of most common poisonous mushrooms such as the sulphur tuft and fly agaric (April–May time frame),[26] but can occur alongside "false morels" (Gyromitra sp.) and "elfin saddles" (Verpa sp.)

Morels in western North America are often found in coniferous forests, including trees in the genera Pinus, Abies, Larix, and Pseudotsuga, as well as in cottonwood riparian forests.[27]

Association with wildfire[edit]

All types of morels may grow abundantly in forests which have been burned by a forest fire,[28] with black morels at the start of the season, followed by the yellows, greys and greens. The mechanism for this behavior is not well known,[15] but appears to be related to both the death of trees and the removal of organic material on the forest floor.[29] Moderate-intensity fires are reported to produce higher abundances of morels than low or high intensities.[17] Where fire suppression is practiced morels often grow in small amounts 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. The Finnish name, huhtasieni, refers to huhta, area cleared for agriculture by the slash and burn method. These spots may be jealously guarded by mushroom pickers, as the mushrooms are a delicacy and sometimes a cash crop.[27]

Efforts to grow morels are rarely successful and the commercial morel industry is based on harvest of wild mushrooms.[27]

False morels[edit]

Main article: False morel

When gathering morels, care must be taken to distinguish them from the poisonous false morels, including Gyromitra esculenta, Verpa bohemica, and others. Although the false morels are sometimes eaten without ill effect, they can cause severe gastrointestinal upset and loss of muscular coordination (including cardiac muscle) if eaten in large quantities or over several days in a row. They contain a gyromitrin-like toxin (an organic, carcinogenic poison) that is produced by the mushroom.

The key differentiating features of false morels in comparison to morels include:[30]

  • The false morels can be told apart from the true morels by careful study of the cap, which is often "wrinkled" or "brainy", rather than honeycomb or net-like. Gyromitra esculenta has a cap that is generally darker and larger than the true morels (Morchella sp.).
  • The caps of early morels (Verpa sp.) are attached only at the apex (top) of the cap, unlike true morels which have caps that are attached at or near the bottom. The easiest way to tell the false from the true variety, is to simply look inside the stem.
  • False morels contain a cotton-ball looking substance inside their stem while true morels are hollow inside.
  • The caps of the false morel can be easily twisted in comparison to the normal morel.
  • False morels are often a brown, reddish color.


Yellow morels in France

Morels are a feature of many cuisines, including Provençal.[31] Their unique flavor is prized by cooks 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 must be cooked before eating. Morels occasionally contain insect larvae that drop out during the drying process.[32]

One of the best and simplest ways to enjoy morels is by gently sauteeing them in butter, cracking pepper on top and sprinkling with salt. Others soak the mushrooms in an egg batter and lightly bread them with saltine crackers or flour before frying them.[citation needed] Lastly, morel mushrooms go great with meat or in a soup.

Morels are not improved by extensive washing or soaking, as it may ruin the delicate flavor and require long cooking times. Due to their natural porosity, morels may contain trace amounts of soil which cannot be washed out. They can best be 'flash frozen' by simply running under cold water or putting them in a bucket to soak for a few minutes, then placing on a cookie sheet or pizza pan and placing into a freezer. After freezing they keep very fresh with the frozen glaze for a long time in airtight plastic containers. However, when thawed they can sometimes turn slightly mushy in the cap.[33] Any visible soil should be removed with a brush, after cutting the body in half lengthwise if needed.

Drying is a popular and effective method of long-term storage for morels, and they are readily available commercially in this form; dried morels can be reconstituted by soaking in warm water or milk. They may also be frozen after steaming or frying. Canning is not recommended because the necessary high pressure and temperature destroys much of the nutty flavor.[citation needed]


Morels contain small amounts of hydrazine[34] toxins that are removed by thorough cooking; morel mushrooms should never be eaten raw.[35] It has been reported that even cooked morels can sometimes cause mild intoxication symptoms when consumed with alcohol.[36]

When eating this mushroom for the first time it is wise to consume a small amount to minimize any allergic reaction. Morels for consumption must be clean and free of decay.

Morels growing in old apple orchards that had been treated with the insecticide lead arsenate may accumulate levels of toxic lead and arsenic that are unhealthy for human consumption.[37]

See also[edit]


  1. ^ "Morchella Dill. ex Pers.". Species Fungorum. CAB International. Retrieved 2014-12-11. 
  2. ^ Segula Masaphy, Limor Zabari, Doron Goldberg, and Gurinaz Jander-Shagug (Spring 2010). "The Complexity of Morchella Systematics: A Case of the Yellow Morel from Israel". Fungi Magazine 3 (2): 14–18. 
  3. ^ Overholts, L.O. (1934). "The morels of Pennsylvania". Proc. Penn. Acad. Sci 8: 108–114. 
  4. ^ Weber, N.S. (1988). In A Morel Hunter's Companion, pp. 111-67. Two Peninsula Press: Lansing.
  5. ^ Korf, R.P. (1973). Discomycetes and Tuberales. In The Fungi (G.C. Ainsworth, F.K. Sparrow, and A.S. Sussman, Eds.), Vol. IVA, pp.249-318. Academic Press: New York.
  6. ^ Kimbrough, J.W. (1970). "Current trends in the classification of discomycetes". Bot. Rev. 36: 91–161. doi:10.1007/bf02858958. 
  7. ^ 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: 762–72. doi:10.2307/3760589. 
  8. ^ Bunyard B.A., Nicholson M.S., Royse D.J. (1995). Phylogenetic resolution of Morchella, Verpa, andDisciotis (Pezizales: Morchellaceae) based on restriction enzyme analysis of the 28S ribosomal RNA gene. Experimental Mycology 19(3):223-33.
  9. ^ Kuo, M. (March 2006). "Morel Data Collection Project: Preliminary results". Retrieved 2009-05-26. 
  10. ^ O'Donnell K, Rooney AP, Mills GL, Kuo M, Weber NS, Rehner SA (Mar 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. 
  11. ^ Harald Kellner, Carsten Renker, and François Buscot (2005). "Species diversity within the Morchella esculenta group (Ascomycota: Morchellaceae) in Germany and France". Organisms, Diversity & Evolution 5 (2): 101–107. doi:10.1016/j.ode.2004.07.001. 
  12. ^ Hatıra Taşkına, Saadet Büyükalacaa, Hasan Hüseyin Doğanb, Stephen A. Rehnerc and 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. 
  13. ^ S. Masaphy, L. Zabari and D. Goldberg (2009). "New long-season ecotype of Morchella rufobrunnea from northern Israel". Micologia Aplicada International 21 (2): 45–55. 
  14. ^ Kanwal HK, Acharya K, Ramesh G, Reddy MS (Dec 25, 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. 
  15. ^ a b c Franck O.P. Stefani, Serge Sokolski, Trish L. Wurtz, Yves Piché, Richard C. Hamelin, J. André Fortin, and Jean A. Bérubé (2010). "Morchella tomentosa: a unique belowground structure and a new clade of morels". Mycologia 102 (5): 1082–1088. doi:10.3852/09-294. PMID 20943507. 
  16. ^ Michael Kuo (July–Sept 2008). "Morchella tomentosa, a new species from western North America, and notes on M. rufobrunnea". Mycotaxon 105: 441–446.  Check date values in: |date= (help)
  17. ^ a b McFarlane, Erika M.; Pilz, David; Weber, Nancy S. (May 2005). "High-elevation gray morels and other Morchella species harvested as non-timber forest products in Idaho and Montana". Mycologist 19 (2): 62–68. doi:10.1017/S0269915X0500203X. 
  18. ^ Kuo M, Dewsbury DR, O'Donnell K, Carter MC, Rehner SA, Moore JD, Moncalvo J-M, Canfield SA, Stephenson SL, Methven AS, Volk TJ. (11 April 2012). "Taxonomic revision of true morels (Morchella) in Canada and the United States". Mycologia 104 (5): 1159–77. doi:10.3852/11-375. PMID 22495449. 
  19. ^ Du, X.-H., Zhao, Q., O’Donnell, K., Rooney, A. P. and Yang, Z. L. (2012). "Multigene molecular phylogenetics reveals true morels (Morchella) are especially species-rich in China". Fungal Genetics and Biology. doi:10.1016/j.fgb.2012.03.006. 
  20. ^ Clowez P. (2012). "Les morilles. Une nouvelle approche mondiale du genre Morchella". Bulletin de la Société Mycologique de France (in French) 126 (3–4): 199–376. 
  21. ^ Richard, Franck; Bellanger, Jean-Michel; Clowez, Philippe; Courtecuisse, Regis; Hansen, Karen; O'Donnell, Kerry; Sauve, Mathieu; Urban, Alexander; Moreau, Pierre-Arthur (30 December 2014). "True morels (Morchella, Pezizales) of Europe and North America: evolutionary relationships inferred from multilocus data and a unified taxonomy". Mycologia (Preliminary version published online). doi:10.3852/14-166. PMID 25550303. 14-166. 
  22. ^ Elliott TF, Bougher NL, O'Donnell K, Trappe JM. (2014). "Morchella australiana sp. nov., an apparent Australian endemic from New South Wales and Victoria". Mycologia 106 (1): 113–8. doi:10.3852/13-065. 
  23. ^ Mattson, D. J.; Podruzny, S. R.; Haroldson, M. A. (2002). "Consumption of fungal sporocarps by Yellowstone grizzly bears". Ursus 13: 95–103. JSTOR 3873191. 
  24. ^ J. L. Dahlstrom, J. E. Smith and N. S. Weber (2000). "Mycorrhiza-like interaction by Morchella with species of the Pinaceae in pure culture synthesis". Mycorrhiza 9 (5): 279–285. doi:10.1007/PL00009992. 
  25. ^ Lincoff, Gary H., The Audubon Society Field Guide to North American Mushrooms (1981) p 326.
  26. ^ Jeanne D. Mihail, Johann N. Bruhn, and Pierluigi Bonello (March 2007). "Spatial and temporal patterns of morel fruiting". Mycological Research 111 (3): 339–346. doi:10.1016/j.mycres.2007.01.007. 
  27. ^ 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. 
  28. ^ Wurtz, Tricia L.; Wiita, Amy L.; Weber, Nancy S.; Pilz, David (2005). Harvesting morels after wildfire in Alaska. Research Note RN-PNW-546. Portland, OR: U.S. Forest Service Pacific Northwest Research Station. 
  29. ^ David F. Greene, Michael Hesketh, and Edith Pounden (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. 
  30. ^ Kuo M. (2007). 100 Edible Mushrooms. Ann Arbor, Michigan: The University of Michigan Press. pp. 55–58. ISBN 0-472-03126-0. 
  31. ^ Olney, Richard (1995). A Provencal Table. London: Pavilion. pp. 31–32. ISBN 1-85793-632-9. 
  32. ^ Wild About Mushrooms: Morels. Mssf.org. Retrieved on 2012-04-17.
  33. ^ Quamut.com, Morel Mushrooms
  34. ^ Paul Stamets Mycelium Running pg 271. (2005)
  35. ^ Ian R. Hall, Peter K. Buchanan (2003). Edible and poisonous mushrooms of the world. Timber Press. ISBN 0-88192-586-1
  36. ^ J. Walton Groves. Poisoning by Morels When Taken with Alcohol. Mycologia, Vol. 56, No. 5 (Sep. - October, 1964), pp. 779-780
  37. ^ Shavit, Elinoar; Shavit, Efrat (Spring 2010). "Lead and Arsenic in Morchella esculenta Fruitbodies Collected in Lead Arsenate Contaminated Apple Orchards in the Northeastern United States: A Preliminary Study". Fungi Magazine 3 (2): 11–18. 
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