Mushrooms of the Pacific Northwest, Revised Edition

LIGHT-SPORED GILLED MUSHROOMS

KEY TO LIGHT-SPORED GILLED MUSHROOM GROUPS

1. Mushroom with more or less distinct cap and stipe; gills thin and plate-like, separate from one another, either attached to stipe to some degree or not attached . . . 2

1. Mushroom often vase- or funnel-shaped without distinct cap and stipe; gills thick and vein- or wrinkle-like, often merging together and/or with cross-veins, extending down stipe . . . Chanterelles (see this page)

1. Mushroom shelf- or bracket-like; stipe lacking or very reduced, sometimes centrally attached but more often attached at the edge of the cap; gills thin and plate-like, extending down stipe when one is present . . . Genus Pleurotus and similar mushrooms (see this page)

2. Gills thick, often widely spaced, and with a waxy appearance; many species small and brightly colored (especially red, orange, or yellow), others larger and duller colored (white, brown, or gray) . . . Wax-Caps (see this page)

2. Gills usually thinner and more closely spaced without a waxy appearance; mushroom size and color varied . . . 3

3. Mushroom usually medium-sized to large; stipe (when present) not particularly slender relative to cap diameter, texture of stipe similar to that of cap . . . 4

3. Mushroom usually small; stipe usually slender relative to cap diameter, texture of stipe often tougher and more pliable than that of cap . . . 9

4. Gills free; stipe often cleanly separable from cap (like a ball-and-socket joint) . . . 5

4. Gills attached to stipe; stipe not cleanly separable from cap . . . 6

5. Young mushroom completely enveloped by a veil, the remnants of which remain as a volva (cup, collar, or rings of tissue) at the base of the stipe and/or a superficial patch or warts on the cap; cap surface (beneath any patch or warts) smooth but outer edge may be grooved . . . Genus Amanita (see this page)

5. Young mushroom not completely enveloped by a veil (but gills may be covered by a partial veil that remains as a ring on the stipe) and no volva, patch, or warts present; cap surface may be smooth, powdery, or innately scaly, outer edge not grooved . . . Genus Lepiota and similar mushrooms (see this page)

6. Texture of cap, gills, and stipe brittle, fresh mushroom may shatter (like automobile safety glass) when thrown against a tree and stipe snapping cleanly like a piece of chalk when broken . . . 7

6. Texture fibrous, not behaving as above . . . 8

7. Fresh mushroom (especially the gills) usually exuding a watery, milky, or colored liquid when cut or broken; color of cap usually dull, color of stipe usually similar to that of cap . . . Genus Lactarius (see this page)

7. Mushroom not exuding liquid when cut or broken; color of cap often bright (red, green, yellow, purple, etc.), color of stipe usually white, sometimes with reddish or purplish blush . . . Genus Russula (see this page)

8. Gills usually decurrent, although sometimes merely attached; cap often vase- or funnel-shaped at maturity; colors usually drab (white, brown, or gray) . . . Genus Clitocybe and similar mushrooms (see this page)

8. Gills attached, often only slightly, but not decurrent; cap usually convex to flat; colors may be dull or fairly bright . . . Genus Tricholoma and similar mushrooms (see this page)

9. Cap usually thin-fleshed and often quite fragile, edge of cap not inrolled when young, cap often conical, at least when young; gills vary from attached to decurrent, those with decurrent gills often exhibit a vase- or funnel-shaped cap; dried specimens not reviving when moistened . . . Genus Mycena and similar mushrooms (see this page)

9. Cap usually somewhat thick-fleshed and tough, edge of cap often inrolled when young, cap usually convex to flat, not conical (although a central hump often is present); gills attached, but not decurrent; dried specimens often reviving when moistened . . . Genus Collybia and similar mushrooms (see this page)

CHANTERELLES

The chanterelles include a variety of mushrooms belonging to five genera (Cantharellus, Craterellus, Gomphus, Polyozellus, and Turbinellus) that are similar in bearing their spores on blunt ridges or veins instead of thin plate-like gills. The fruitbodies range from small to large and have a cap and stipe, although there is not always a clear distinction between them; in many cases, they are more or less vase-, funnel-, or trumpet-shaped with the fertile ridges extending down the stipe. They are brittle, fleshy, or somewhat leathery in texture, never woody. The stipe can be solid or have a hollow center. The fertile surface often has a waxy luster and comprises an anastomosing system of thick, shallow folds with blunt edges and cross-veins; it is nearly smooth in some species of Craterellus and Cantharellus, but only one of these, Craterellus calicornucopioides, occurs in the PNW. The chanterelles are strictly woodland fungi, as they are ectomycorrhizal with a variety of trees. Thus, they are found on soil or sometimes on well-rotted wood.

The chanterelles are not closely related to the gilled mushrooms and, despite their similar appearance, the different chanterelles are not all closely related to one another. Cantharellus and Craterellus indeed are closely related to each other, as well as to Hydnum (the hedgehogs). On the other hand, Gomphus and Turbinellus are closely related to coral-fungi such as ramarias, stinkhorns (and their truffle-like relatives), and most earthstars. Polyozellus is most closely related to the thelephoras and spine-fungi in the genera Hydnellum and Sarcodon.

Cantharellus formosus Corner

Pacific golden chanterelle

Although nearly all the golden chanterelles in North America have been called Cantharellus cibarius, ongoing studies have confirmed that they actually represent many different, but similar-looking, species. The species epithet formosus (“finely formed,” “beautiful”) is certainly descriptive of many of our golden chanterelles. The fruitbodies are often large for a chanterelle, and have a dull orange to brownish orange cap that readily bruises brownish and often is finely scaly. The fertile ridges often are deep and relatively thin; they are usually pale orange-yellow but may have a pinkish cast. The stipe usually is fairly slender and tapered downward. Cantharellus formosus fruits abundantly throughout moist portions of the PNW and is the species most commonly found for sale in produce markets and grocery stores, as well as on restaurant menus.

Gilled mushrooms that could be confused with golden chanterelles include Hygrophoropsis aurantiaca, Chroogomphus tomentosus, and, in California, Omphalotus olivascens. All three have sharp, blade-like gills (although those of H. aurantiaca can be deceivingly chanterelle-like), none of these is worth eating, and O. olivascens is poisonous. Turbinellus floccosus and T. kauffmanii both have blunt fold-like gills like chanterelles; however, both are generally more vase-like and have coarse scales on the cap. Older, orange-stained individuals of the white chanterelle (Cantharellus subalbidus) and individuals of other golden chanterelle species such as C. cascadensis can be hard to differentiate from Pacific golden chanterelles, but all are good edibles.

Cantharellus cascadensis Dunham, O’Dell & R. Molina

Both the golden chanterelle Cantharellus cascadensis and C. roseocanus (Redhead, Norvell & Danell) Redhead, Norvell & Moncalvo (rainbow chanterelle) first passed as C. cibarius, and then as C. formosus, but now are recognized as separate entities in large part as a result of molecular analyses. They are very similar to C. formosus in most respects, differing primarily in sometimes subtle details of stature, coloration, bruising reaction, scaliness, and habitat. The cap of C. cascadensis usually exhibits bright orange-yellow hues and a smooth or slightly woolly surface. The stipe often is club-shaped or bulbous. The details of its distribution are not yet known, although it appears to be most common in the Oregon Cascades, extending southward into California. The cap of C. roseocanus is smooth, bright yellow-orange, often has a pinkish blush near its edge, and bruises less than the other two golden chanterelles. It is associated primarily with spruce, occurring with Sitka spruce near the coast and with Engelmann spruce in the mountains, but also has occasionally been reported from forests lacking spruce. Cantharellus formosus and C. cascadensis appear to be restricted to the PNW and northern California, whereas C. roseocanus is thought to occur more widely, including in the southern Rocky Mountains of Colorado and New Mexico.

Cantharellus subalbidus A.H. Smith & Morse

white chanterelle

Cantharellus subalbidus differs from the golden chanterelles primarily by its cream to ivory color. It darkens to yellow-orange with age and so older specimens sometimes can be difficult to distinguish from golden chanterelles, especially C. cascadensis, which is similarly stocky. Although it occurs in a variety of forests containing Douglas-fir and hemlock, it seems to have a greater affinity for old forests than do C. formosus and C. cascadensis.

Craterellus tubaeformis (Fries) Quélet

winter chanterelle, yellow foot

Craterellus tubaeformis is a small, slender, trumpet-shaped chanterelle with a brownish or orange-brown cap, hollow stipe, and penchant for growing on mossy rotten wood. It has a long fruiting season although, in most of the PNW, it is not common in winter (it is in California, though). Previously it was known as Cantharellus tubaeformis and also as Cantharellus/Craterellus infundibuliformis. Results of the molecular analysis that supported Craterellus as the appropriate genus for C. tubaeformis also suggest that the winter chanterelle of the PNW is not that species. Thus, we need a new name for our fungus, and C. neotubaeformis has been suggested, although not yet formally proposed. Despite its size, C. tubaeformis is edible and considered choice by some. Its tendency to grow in large troops allows it to be gathered in sufficient quantity to be worthwhile, including by commerical pickers.

Gomphus clavatus (Persoon) Gray

pig’s ear

According to Alexander Smith, “This is the fungus usually referred to when someone asks about ‘that funny-looking thing which is purplish underneath.’ . . . It is a most peculiar fungus to say the least.” The purplish, veined fertile surface combined with the flat to funnel-like tan cap and growth in clusters make it distinctive. The Polyozellus species also are usually clustered and have a veined fertile surface, but are entirely dark blue, purplish, or black. Gomphus clavatus is less abundant than many mycophagists would like, but is not rare, occurring throughout the conifer forests of western and northern North America. It is considered a choice edible by some and less desirable by others but, unlike the other chanterelles, it often is insect-infested.

Genus Polyozellus

blue chanterelle, black chanterelle

The dark bluish, purplish, or blackish color, veined fertile surface, and tendency to grow in tight-packed clusters make the stipitate Polyozellus species easy to identify as a group. If confirmation is necessary, the warty white spores and blackish green reaction of the flesh when potassium hydroxide is applied (seen most easily in thin section under the microscope) will clinch the identification. Recently, it has been shown that, rather than a single worldwide species, P. multiplex, there are at least five species in the complex, including three that occur in the PNW. All three of our species are relatively uncommon and tend to occur most frequently in old-growth forests. Polyozellus atrolazulinus S.A. Trudell & Kõljalg is the easiest of the three to identify—its spores are less than 7 μm in diameter, the cap flesh usually thinner and darker gray, and the overall color more likely to be obviously bright bluish, at least when young, compared to P. marymargaretae Beug & I. Saar (named for Mary Margaret “Maggie” Rogers) and P. purpureoniger Spirin & I. Saar. The latter two species are more difficult to tell apart as both have spores typically 7–8 μm in diameter and relatively thicker and lighter gray cap flesh. Polyozellus marymargaretae tends to be bluish when young whereas P. purpureoniger is more purplish but both become black in age. In the PNW, P. atrolazulinus seems to be the most widespread of the three species and P. marymargaretae the most common, especially in Oregon and southern to central Washington. Polyozellus purpureoniger seems to be the least common and to have a more northerly range (Washington–B.C.–southern Alaska), although at least one collection has been made in southern Oregon. Some consider the blue/black chanterelles to be good edible species, while others are much less enthusiastic about them. They also are sought after by dyers as they can impart blue-green colors to wool and silk.

Turbinellus floccosus (Schweinitz) Earle ex Giachini & Castellano

woolly chanterelle

Not truly woolly, Turbinellus floccosus is more accurately characterized by the coarse scales that usually line its deep vase-shaped cap; however, it is a highly variable fungus, and the degree of scaliness is by no means constant. In its common form, the cap when fresh is a deep reddish orange but fades with age, and old pale specimens can be found that might seem to be a different species. The fertile surface is whitish to pale yellowish and highly wrinkled and forked, with portions appearing almost like pores in older specimens. The size of the fruitbodies varies from small-medium to fairly large, and the shape can be tall and slender or short and squatty with the vase shape being more or less developed. It is common throughout the conifer forests of western and northern North America. Gomphus bonarii has been said to differ by being smaller, having block-like yellow-orange scales with red tips, whitish hymenium with no traces of yellow or orange, and a tendency to grow in clusters. However, in practice, it is very difficult to distinguish two species, and the most recent critical study did not recognize G. bonarii as a separate species, although some mycologists disagree. Although some people consider T. floccosus delicious, others find the taste poor, and still others have reported gastric discomfort after eating it.

GENUS RUSSULA

Russula is a particularly easy genus to recognize, and some of its species also can be identified readily. However, most species are difficult to identify both because of the nature of variation within the genus and the species and because its taxonomy in North America is currently a nightmare. Most russulas are medium-sized to large woodland mushrooms with colorful caps, white stipes, and a characteristic squatty appearance resulting from the width of the caps being about the same as the height of the often thickish stipes. They are very clean-looking in part because they lack veils; thus, they do not have rings or volvas (at least the species in North America do not). The other distinctive characteristic is their brittle texture—a fresh russula thrown against a tree will shatter like automobile safety glass into small smooth-edged chunks (I recommend that everyone try this once, but not make a habit of destroying russulas as, when intact and in place, they are a very attractive visual element in our forests). Other than in Lactarius and other close relatives of russulas, in which it usually is less well developed, this brittle texture is very rare in mushrooms. A less violent means of experiencing it is to break a fresh stipe in half—it will break cleanly and audibly, like a piece of chalk. The reason for this texture can be seen under the microscope—the flesh includes nests of spherical cells that look much like soap suds in a sink of dishwater. When shearing force is applied, these cells can move fairly freely past one another, unlike long slender hyphae that are entangled like a mass of spaghetti. Another characteristic microscopic feature is the spores, which are ellipsoid to nearly globose, with warts or ridges or networks of ridges that stain bluish black in Melzer’s reagent. In mass, the spores vary from pure white, through shades of cream and yellow, to fairly dark ocher. The flesh of many species is acrid either immediately or after a minute or so, and it has led some mushroomers to refer to “russula mouth.”

The members of Russula and Lactarius are closely related, usually being separated macroscopically by the presence of latex in Lactarius. In addition, the more brittle flesh of the russulas, their tendency to have more brightly colored caps, and their more consistently squatty stature make recognition of the two genera fairly easy in most cases. In recent years, molecular and morphological studies have shown that the russulas and lactariuses belong to an evolutionary line distinct from that of other gilled mushrooms. The molecular analyses also indicate that Russula and Lactarius, as traditionally configured, do not represent natural evolutionary groups. Instead many species of “Lactarius” cluster with the russulas. In light of this, a new genus, Multifurca, was created and an old genus, Lactifluus, was resurrected so that there are currently four genera where we were used to having two. However, so far, our PNW species all fit within the traditional Russula versus Lactarius arrangement. Both russulas and lactariuses are ectomycorrhizal, even the species that can be found fruiting on rotting wood, and hence are found in forests and other areas where suitable host trees are present.

It is not known how many species of Russula we have in the PNW, however some estimates suggest a few hundred, most of which are unnamed or otherwise poorly known. For instance, the late Ben Woo, the PNW’s most ardent student of russulas, made over 1,000 well-documented collections that were studied by Dr. Anna Bazzicalupo. Ben’s collections conservatively represented 72 different species, to which initially only 28 could be assigned a confident name. Some of the new species that resulted from that study and follow-up work with Danny Miller are included in this section.

As for edibility of russulas, Ben said it best: “Russulas are so abundant that the question of their edibility is always raised by beginners. The answer is usually a wry, ‘Well, if they taste mild, you can try eating them.’ The implication is that they are not particularly choice. It’s the plain truth that, given the availability of almost any other edible mushrooms, russulas come off a distant second. Most are either of poor quality or unpalatable.” The only species he recommended are Russula olivacea and R. xerampelina (both sensu PNW, as we have very close relatives of both, but probably not the actual European species). However, other than reports of fatal poisonings in Japan by R. sub-nigricans, russulas are generally considered safe to eat if not necessarily choice.

Russula adusta (Persoon) Fries

Russula adusta is a member of the subgenus Compactae, which includes relatively large, dense, hard-fleshed mushrooms that are white to brownish and often blacken in age, sometimes after an intermediate stage of reddening. Russula adusta blackens only slightly, and the flesh pinkens lightly when cut. The viscid, then relatively shiny, cap is brownish to grayish (adustus is Latin for “burned” or “tanned”), and the odor is said to approximate that of empty wine barrels (although this odor has not always been noticed in PNW collections). The spores are white and the taste is mild. It grows with conifers. The other blackening russulas in the PNW differ by having dry to only subviscid caps. The most similar of these is what has been called R. densifolia in the PNW. It has a more yellowish brown cap, reddening flesh, perhaps a more acrid taste, and perhaps slightly yellowish spores. Russula albonigra is whiter at first, and then turns black without reddening, with the gills often the last part to blacken. Ongoing molecular studies suggest that in the PNW we have something possibly the same as the “real” R. adusta as well as two very close relatives that Danny Miller refers to as aff. adusta #1 and aff. adusta #2.

Russula aeruginea Lindblad ex Fries

Russula aeruginea is a common species often found with spruce. It has a smooth, shiny gray-green or green cap (hence the species epithet from Latin aeruginis, meaning “verdigris” or “copper rust”) that often develops rusty spots, cream gills that may also become rusty spotted, white stipe that can develop brownish spots near the base, a mild or very slowly and slightly acrid taste, and deep cream spore-print. The spores are broadly ellipsoid, 6–9 χ 5–7 μm, and ornamented with mostly isolated warts and short ridges. Molecular analyses indicate that our species is at least a very close relative of the European species, which is associated with birch and spruce, and might be the same.

Russula benwooii Bazzicalupo, D. Miller & Buyck

Russula benwooii is a prime example of why russula identification can be extremely frustrating. Its fruitbodies range from medium-sized to large (they can be among the largest in the genus), the cap is viscid when humidity is high, spore-print and mature gills cream to pale yellow, stipe white and sometimes flushed purple or grayish lilac, and the taste is mild. The potential for frustration arises from the extreme variability of the cap color. It is most often brown to brownish tan, sometimes mixed with olive-green, reddish pink, wine-red, or purple. However, it can also be entirely wine-red, olive-green, or purple, as in the photograph. The center of the cap can be the same as, or lighter or darker than, the remainder. Russula benwooii is a very common species, associated with western hemlock and perhaps other conifers. It differs from the similarly variable R. xerampelina