|Morchella elata clade|
|Morchella esculenta clade|
Pamela Kaminski (e. N.Am.)
NEW ARTICLE December 2014
Richard, Franck, Mathieu Sauve, Jean-Michel Bellanger, Philippe Clowez, Karen Hansen, Kerry O'Donnell, Alexander Urban, Régis Courtecuisse, and Pierre-Arthur Morea. 2014. True Morels (Morchella, Pezizales) of Europe and North America: Evolutionary relationships inferred from multilocus data and a unified taxonomy. Mycologia: preliminary version published on December 30, 2014 as doi:10.3852/14-166.
This article on morels with proposed name changes will be published in Mycologia by Richard et al. It resolves some of the name conflicts between the 2012 publications below. The authors of Kuo et al. article appear to have been unaware of the publication of new names earlier in the year by Clowez.
Kuo, Michael, Damon R. Dewsbury, Kerry O'Donnell, M. Carol Carter, Stephen A. Rehner, John David Moore, Jean-Marc Moncalvo, Stephen A. Canfield, Stephen L. Stephenson, Andrew S. Methven, Thomas J. Volk. 2012. Taxonomic revision of true morels (Morchella) in Canada and the United States. Mycologia 104(5): 1159-1171.
Clowez, Phillippe. 2010. Les morilles, une nouvelle approche mondiale du genre Morchella. Bull. soc. mycol. France 126(3-4):199-376.
Current species established for the Pacific Northwest following the Richard et al. Mycologia article are the following. Characters are mnemonic and not meant to characterize the species completely.
|Current Name||Kuo et al.||Clowez||O’Donnell et al.||English name|
|M. tomentosa||M. tomentosa||M. tomentosa||Mel-1||fuzzy-foot morel|
|M. tridentina||M. frustrata||M. elatoides||Mel-2||blond morel|
|M. populiphila||M. populiphila||Not covered||Mel-5||western half-free morel|
|M. sextelata||M. sextelata||Not covered||Mel-6||a burn morel|
|M. eximia||M. septimelata||M. eximia
|Mel-7||a burn morel|
|M. exuberans||M. capitata||M. exuberans||Mel-9||a burn morel|
|M. importuna||M. importuna||M. elata
|M. snyderi||M. snyderi||Not covered||Mel-12||Snyder’s morel|
|M. brunnea||M. brunnea||Not covered||Mel-22||natural black morel|
|M. americana||M. esculentoides||M. americana
|Mes-4||American. yellow morel|
|M. prava||M. prava||Not covered||Mes-7|
|M. rufobrunnea||M. rufobrunnea||M. rufobrunnea||Mrb||blushing morel|
(Pink and green morels were found in more than one 'burn black' proposed species. Gray morels are often black fuzzyfoots.)
The morels and false morels have some similarity in general appearance. Fruitbodies normally have a distinct stipe (stem) and a head (cap) that is usually convoluted in some way (or saddle-shaped in the case of some false morels). Spores are borne on the outer surface (or in large pits in the outer surface). Presently they fall into two families, Helvellaceae and Morchellaceae.
The Trial Key to HELVELLACEAE in the Pacific Northwest was written by Harold Treibs and covers Discina, Gyromitra, and Helvella.
There is no comprehensive key to Morchellaceae for the good reason that the taxonomy is partly based on recent molecular research and the macroscopic appearances, ecology, and distribution are still being worked out. It is easy to distinguish the thimble morels (Verpa conica and Verpa bohemica) from true morels (Morchella).
Molecular techniques have been used in an attempt to untangle species concepts. This account will discuss Jung(1) 1993, Bunyard(1) 1994, Pilz(2) 2004, Kuo(2) 2006, Clowez(1) 2010, O'Donnell(1) 2011, Kuo(6) 2012, and Richard(1) 2014.
Early molecular research suggested that the black morels (known by the names M. angusticeps, M. elata, M. conica) and the yellow morels (known by the names M. esculenta, M. crassipes, and M. deliciosa) are separate taxonomic groups (Bunyard), and immunological techniques suggested the gray, tan, and large tan forms in the M. esculenta complex are immunologically indistinguishable and likely conspecific. (Jung).
Five putative morel species were designated with the aid of molecular techniques: PS A, PS B, PS C, PS D, and PS E. They had field characters as follows. PS A is a member of the black morel grouping, called the natural black morel because it occurs away from fires: it has mature pits grayish tan or pale tan. PS B is a second member of the black morel grouping and occurs after fires. It includes the so called pink morel with mature pits rosy tan to pinkish brown. PS C is a third member of the black morel grouping, and also occurs after fires. It includes the green morel (so called because of the olive tinged gray or olive tinged brown color of mature pits). These three black morels all key out to the M. elata (complex), M. conica, or M. angusticeps, depending on the reference used, but all of these names are of uncertain application. PS D is the black stocking morel, a common member of the "gray morel" grouping. It has the surface of the stem distinctly velvety when young, dark brown to black (as opposed to color at all ages entirely off-white, ivory, tan, dull pink or dull purple), becoming paler when old "as the velvety layer is stretched apart (use a hand lens to look for dark tufts of hyphae in old specimens)". This PS D morel is one of the burn morels, fruiting the spring following a fire. It had been called Morchella atrotomentosa (see for example McKnight 1987), but there are nomenclatural problems with the use of that name. Later it was described by Kuo 2008 as Morchella tomentosa Kuo. PS E (mountain blond morel) occurs away from fires and has the ribs separating the pits lighter in color (ivory to nearly white) than the lining of the pits when old. By maturity the ribs are off-white sometimes with amber stains and bruises. Pits are gray when very young, becoming tan, golden, or straw yellow when mature. It differs from most descriptions of M. esculenta from Europe or elsewhere in North America in that 1) it has a relatively narrow head, rather than oval or rounded, especially in young specimens, 2) "the primary ribs are strongly vertical and relatively straight producing elongated pits rather than the rounded to somewhat irregular pits generally attributed to M. esculenta", and 3) it "seems to be characteristic of conifer forests including either lodgepole or ponderosa pine, whereas the complex centered on M. esculenta is commonly associated with hardwoods sometimes mixed with conifers." Both PS E and a morel that fits the classic concept of M. esculenta occur in eastern Oregon. (Pilz 2004).
The Morel Data Collection Project, outlined in Kuo(2005) and Kuo(2006), included a number of Pacific Northwest taxa:
Two years later Mycologia released a pre-publication version of an article that applied Latin names to these taxa. Unfortunately, and apparently unbeknownst to the authors, a French monograph of Morchella worldwide by Clowez had already described a number of North American species. Four new species that appear to apply to the Pacific Northwest were
English names applied to Morchella tomentosa include "black stocking morel", "black foot morel" and "fuzzy-foot". It has been called a "gray" morel, but that term is applied by different people to different morels. The term "gray" is sometimes used of yellow morels at a certain stage of development or under certain ecological conditions. The term "yellow morel" includes the morels resembling Morchella esculenta of Europe in colour, and has been applied to the esculenta clade. The term "black morel" has been applied to the whole elata clade but traditionally comprises those with prominently dark colors including the natural black morel, the green morel, the pink morel, and sometimes the black stocking morel. Green burn morels and pink burn morels are however found in more than one of the taxa of the elata clade designated in O'Donnell 2011 and Kuo 2012.
Black morels develop in burn areas earlier in the season than the gray and green members of the black burn morel group. They have light stems that fade to rusty brown, and the ridges turn black. There is often a taproot. (Larry Evans, speaking of Montana in particular, pers. comm.)
Richard 2014 synonymized
a) M. frustrata and M. elotoides with M. tridentina,
b) M. septimelata, M. anthracophila, and M. carbonaria with M. eximia
c) M. capitata with M. exuberans, and
d) M. esculentoides and M. californica with M. americana.
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