MORCHELLACEAE in the Pacific Northwest

An Introduction



by Ian Gibson, South Vancouver Island Mycological Society

Copyright © Pacific Northwest Key Council, 2007, 2009, 2012, 2015, 2019

Photo copyright held by each photographer

Do not copy photos without permission

Morchella elata clade



Steve Trudell Morchella esculenta clade



Pamela Kaminski (e. N.Am.)

The most important three articles for our purpose in the last few years are Clowez (2010), Kuo et al. (2012) and Richard (2015). The third made an attempt to co-ordinate and expand on the findings of the first two. Morchella eohespera was described in 2016. 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. 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 104(5): 359-382.

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.

Morchella tomentosa - "black fuzzyfoot" burn morel

- "black fuzzyfoot" burn morel Morchella tridentina - blond morel colors but black morel shape, ridges do not darken with age, forests, (includes what is referred to as 'mountain blond morel' or 'western blond morel')

- blond morel colors but black morel shape, ridges do not darken with age, forests, (includes what is referred to as 'mountain blond morel' or 'western blond morel') Morchella populiphila - western North American half-free morel, associated with Populus trichocarpa

- western North American half-free morel, associated with Morchella sextelata - a 'burn black', microscopic elements on ridges cylindric, subclavate, subfusiform, or subcapitate, distinguished from M. eximia only by DNA

- a 'burn black', microscopic elements on ridges cylindric, subclavate, subfusiform, or subcapitate, distinguished from only by DNA Morchella eximia - a 'burn black' that includes M. anthracophila , M. carbonaria , M. septimelata

- a 'burn black' that includes , , Morchella exuberans - a 'burn black', includes Morchella capitata with microscopic elements on ridges overwhelmingly capitate

- a 'burn black', includes with microscopic elements on ridges overwhelmingly capitate Morchella importuna - a 'black' morel found in wood-chip and garden/landscape settings, cross ridges often give a ladder-like appearance, often several in clusters

- a 'black' morel found in wood-chip and garden/landscape settings, cross ridges often give a ladder-like appearance, often several in clusters Morchella snyderi - pale to greenish to burgundy when young but black morel form, ridges darken with age, stipe often with conspicuous lacunae/ridges (even when young) and with granularity, typically in clusters of 2-20, generally non-burn with montane conifers

- pale to greenish to burgundy when young but black morel form, ridges darken with age, stipe often with conspicuous lacunae/ridges (even when young) and with granularity, typically in clusters of 2-20, generally non-burn with montane conifers Morchella 'Mel-19' - unnamed but described in 2016 as Morchella eohespera , similar to Morchella brunnea but with conifers

- unnamed but described in 2016 as , similar to but with conifers Morchella brunnea - 'natural black', grows singly in forest with hardwoods

- 'natural black', grows singly in forest with hardwoods Morchella americana - North American 'esculenta' yellow morel, with hardwoods

- North American 'esculenta' yellow morel, with hardwoods Morchella prava - like a Morchella americana with contorted ridges and pits

- like a with contorted ridges and pits Morchella rufobrunnea - pale ridges, dark pits, yellow morel form, developing orangish to reddish stains, wood-chip and landscape settings

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

M. anthracophila

M. carbonaria Mel-7 a burn morel M. exuberans M. capitata M. exuberans Mel-9 a burn morel M. importuna M. importuna M. elata

M. vaporaria Mel-10 landscape morel M. snyderi M. snyderi Not covered Mel-12 Snyders morel M. eohespera Unnamed Unnamed Mel-19 M. brunnea M. brunnea Not covered Mel-22 natural black morel M. americana M. esculentoides M. americana

M. californica Mes-4 American. yellow morel M. prava M. prava Not covered Mes-7 M. rufobrunnea M. rufobrunnea M. rufobrunnea Mrb blushing morel

MORELS AND FALSE MORELS

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 and DISCINACEAE in the Pacific Northwest was written by Harold Treibs and covers Discina, Gyromitra, and Helvella.

HISTORY

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:

Morchella rufobrunnea, the fuzzy-foot morel which was later named Morchella tomentosa, several taxa similar to the natural black morel in appearance but distinguished by DNA methods, three to five according to the method used, and all recorded (among other places) in Oregon. a half-free morel, the western blond (mountain blond renamed to include not only mountain conifer habitat but lowland hardwood habitat), and a taxon resembling the European Morchella esculenta.

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

Morchella anthracophila, a black burn morel described from British Columbia, Morchella carbonaria, another black burn morel described from British Columbia, Morchella fallax, which corresponds to the so-called western blond, and Morchella americana, the common yellow morel that went in years past under the European name Morchella esculenta, often identical in color but with the cap more elongated, and the pits more numerous, slightly long, smaller, and less regular.

Morchella rufobrunnea (pale ridges, dark pits, yellow morel form, developing orangish to reddish stains, wood-chip and landscape settings), Morchella tomentosa (black stocking burn morel).

Morchella capitata, M. sextelata, and M. septimelata, three burn black morels, the first perhaps distinguished by a predominance of microscopic capitate elements on the ridges, Morchella brunnea, a "natural black" occurring in non-burned forests, associated with hardwoods, probably also to be expected with non-burned conifers Morchella importuna (a black morel with a laddered pattern of pits/ridges, favoring wood-chip and garden/landscape settings), Morchella populiphila (western North American half-free morel associated with Populus trichocarpa), Morchella snyderi (pale colors when young but black morel form, ridges become brown, stem often with conspicuous pits/ridges (even when young) and with granularity, occurring under montane conifers), Morchella frustrata (pale colors but black morel form, in forests, likely what has been called the mountain blond or western blond, and may be the same as Morchella fallax), and Morchella esculentoides (the North American esculenta-like yellow morel that is likely the same as Morchella americana).

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.



REFERENCES

Arora, David. 1986 Mushrooms Demystified Second Edition. Ten Speed Press, Berkeley. Breitenbach, J., Kränzlin, F. 1984. Fungi of Switzerland Volume 1 Ascomycetes. Edition Mykologia Lucerne. Bunyard, Britt A., Michael S. Nicholson, Daniel J. Royse. 1994. "A systematic assessment of Morchella using RFLP analysis of the 28S ribosomal RNA gene." Mycologia 86(6): 762-772. Clowez, Phillippe. 2010. "Les morilles, une nouvelle approche mondiale du genre Morchella." Bull. soc. mycol. France 126(3-4):199-376 Jung, C.H. 1993. "Systematics of Morchella esculenta complex using enzyme-linked immunosorbent assay. Mycologia 85(4): 677-684. Kuo, Michael. 2005. Morels. University of Michigan Press. Kuo, Michael. 2006, March. North American Morels in the MDCP. Retrieved from the MushroomExpert.Com Web site: http://www.mushroomexpert.com/mdcp/results_legent.html Kuo, Michael. 2008. "Morchella tomentosa, a new species from western North America, and notes on M. rufobrunnea." Mycotaxon 105: 441-446. 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, Androw S. Methven, Thomas J. Volk. 2012. Taxonomic revision of true morels (Morchella) in Canada and the United States. Mycologia 104(5): 1159-1177. McFarlane, Erika M., David Pilz, and Nancy S. Weber. 2005. "High-elevation gray morels and other Morchella species harvested as non-timber forest products in Idaho and Montana." Mycologist 19: 62-68. McKnight, Kent H., McKnight, Vera B. 1987. A Field Guide to Mushrooms North America. Peterson Field Guide Series. Houghton Mifflin, New York. O'Donnell, Kerry, Alejandro P. Rooney, Gary L. Mills, Michael Kuo, Nancy Weber, Stephen A. Rehner. 2011. "Phylogeny and historical biogeography of true morels (Morchella) reveals an early Cretaceous origin and high continental endemism and provincialism in the Holoarctic." Fungal Genetics and Biology 48: 252-265. Pilz, David, Nancy S. Weber, M. Carol Carter, Catherine Parks, and Randy Molina. 2004. "Productivity and diversity of morel mushrooms in healthy, burned, and insect-damaged forests of northeastern Oregon." Forest Ecology and Management 198: 367-386. 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: 104(5): 359-382. Tylutki, Edmund E. 1979. Mushrooms of Idaho and the Pacific Northwest. Volume 1. Discomycetes. University of Idaho Press, Moscow, Idaho. Voitk, Andrus, Michael W. Beug, Kerry ODonnell & Michael Burzynski. 2016. Two new species of true morels from Newfoundland and Labrador: cosmopolitan Morchella eohespera and parochial M. laurentiana. Mycologia 108(1): 31-37. Weber, Nancy Smith. 1988. A Morel Hunters Companion A Guide to the True and False Morels of Michigan. Chief Photographer James A. Weber.

- END -