Key to genera of Toothed Fungi
These notes are designed to be helpful in identifying seven genera of stipitate (stemmed) toothed fungi as they occur in the Pacific Northwest, with comments on the most commonly found species in each genus. Hericium is usually included with this group, although it does not have a true stem. Dr. Kenneth Harrison, the North American expert, says of the group, "The remarkable longevity of individual sporophores of many species and the changes in appearance that occur during the long period of their development have confused all workers studying this group." Amen!
The fruitbody can be fleshy (firm or fragile), leathery or woody, (gelatinous only for Pseudohydnum gelatinosum). Spores can be brown or white, rough or smooth – but all are borne on projecting or downward hanging spines or teeth. Often a pleasant fragrance remains after drying.
Six genera have white spores: Auriscalpium, Hericium, Hydnum, Phellodon, Pseudohydnum, and Bankera. These genera have only a few species in the Pacific Northwest and their species descriptions are given here. Following the molecular findings of Niskanen et al.(2018), at least six species of Hydnum have been found in the Pacific Northwest. See the note below under Hydnum later in this Introduction.
Two genera have brown spores: Sarcodon and Hydnellum. They are considered in detail in separate documents, but the borderline between the two has been obscured by the molecular findings appear to include most species known as Sarcodon in the same clade as Hydnellum. See the note below under Sarcodon.
Some confusion could result from the fact that the Sarcodon species were formerly called Hydnum and the Hydnum species formerly called Dentinum. The older names are used in the excellent field guide Mushrooms Demystified by David Arora.
Traditionally Hydnum, Hericium, Phellodon, Bankera, Sarcodon, and Hydnellum are studied together, and they have certain similarities that once placed them all in the family Hydnaceae. All but Hydnum have been moved, Hericium to Hericiaceae, and the rest to Bankeraceae. Baird et al.(2013) synonymized Bankera with Phellodon. The process of determining genetic relationships by modern molecular techniques is not yet complete.
Auriscalpium vulgare is included here because of its obvious teeth. Pseudohydnum gelatinosum, the so-called toothed jelly, in unrelated and quite different, but is very common and does have teeth on the underside of its cap. (This would presumably be a case of convergent evolution: there are only so many ways that the spore-bearing area can increase its surface.) The term "stipitate" excludes certain polypores that have teeth on the underside, notably Echinodontium tinctorium, and some fruitbodies of others like Trichaptum abietinum may show tooth-like patterns on the underside. An array of polypores and crust fungi that grow mostly flat on wood show a toothed surface (Hyphodontia for instance), but these are not considered further here as their identification would normally require microscopic examination.
Auriscalpium – Fruitbody small (2-7 cm tall), tough, brown hairy cap and lateral stipe, growing only on decaying buried conifers cones – in most of northern U.S. on pine but in this area on Douglas-fir. A single species, Auriscalpium vulgare.
Hericium – Fruitbody is a framework of branches with projecting or hanging spines of various sizes, loose or compact. All grow on wood and are edible. Four species occur in the PNW.
H. erinaceus – solid mass with spines that can be 6 cm long, white, yellowing in age; grows only on deciduous wood, oak along Pacific Coast, in this area on maple; common name Bear’s Head.
H. coralloides – branches intricate and fused, spines short, pure white; grows only on deciduous wood, usually poplar.
H. abietis – variable in size but can be massive when mature, up to 75 x 25 cm, compact or openly branched, white to cream when young, yellowish gray or darker in age; growing on conifers; several forms include forma brevispineum with short spines (1-3 mm) on flexuous branches and forma weirii, very compact with short branches hidden by spines (20-25 mm).
H. americanum – similar to H. abietis but growing on hardwoods, spore size different.
Hydnum – Fruit body fleshy, brittle; stipe central, eccentric or lateral; in moss or duff under mixed conifers. Two species, both edible have long been recognized in the Pacific Northwest. Niskanen et al.(2018) described more species, so that there is evidence of at least six species. We include in the Hydnum repandum group Hydnum neorepandum, Hydnum olympicum, and Hydnum melleopallidum (with fruitbodies that have predominantly decurrent teeth and predominantly pale pinkish orange to pale brownish caps, and may be larger and more robust ). We include in the Hydnum umbilicatum group Hydnum oregonensis, Hydnum cf. umbilicatum, and Hydnum melitosarx (with fruitbodies that have predominantly non-decurrent teeth, and may have predominantly ochraceous to orange brown caps).
H. repandum – cap 3-10 cm broad, convex to slightly depressed, yellowish white to pale orange becoming darker when bruised; stipe short, thick.
H. umbilicatum – smaller, abruptly umbilicate, deeper orangey color; stipe thin and long.
Phellodon – Fruit body small to medium size; thin cap, often zoned; flesh tough, fibrous; spines pale gray to buff; singly or in fused masses under mixed conifers. Three species, possibly more.
Phellodon tomentosus – some shade of brown, strongly zoned, margin whitish; up to 6 cm tall, 5.5 cm broad; fragrant.
Phellodon melaleucus and P. atratus – very similar with dark gray blue to black coloration; the former is velvety, more delicate, with a thin radicating stipe; the latter has whitish margin, is more robust, very abundant in second growth conifer.
Bankera – Fruit body large (8-15 cm), some shade of brown with brittle flesh and whitish spines. (According to K. Harrison this genus bears the same relationship to Phellodon in white-spored group as Sarcodon does to Hydnellum in brown-spored group.) [Both species have proposed current names in Phellodon.]
Bankera fuligineoalba – large dark brown cap with wavy lighter margin; short stout mushroom often covered with duff. (One collection reported from Oregon.)
Bankera violascens – grayish brown cap becoming cracked and scaly with split, incurved margin.
Sarcodon – The cap is fleshy, sometimes fragile, like a conventional gilled mushroom. 14 species or more may occur in PNW; collections often do not fit descriptions. Among the more recognizable are:
S. imbricatus – large with large medium brown upturned scales on lighter brown background, often sunken disk cap, preferred Sitka spruce forest habitat, and dyeing wool. grayish beige
S. squamosus – separated from S. imbricatus by vinaceous to brown cap, often dark vinaceous brown to purplish black, slightly smaller cap scales, lesser likelihood of a deeply sunken disk, preferred pine habitat, and dying wool bluish green not grayish beige. Older guides include it as part of S. imbricatus.
S. scabrosus – with smaller flattened scales; long (1 cm) spines interspersed with short ones; taste bitter, stem base olive-black.
S. fuscoindicus – gray-lavender cap and stipe, with firm purple flesh.
S. stereosarcinon – large, dark reddish brown, zoned; drying hard and woody – like a cross with Hydnellum.
S. calvatus – very large (15-28 cm) cream buff cap; variety odoratum has strong sweet fragrance.
S. atroviridis – grayish tan with enlarged stem base; entire mushroom olive green when dry.
Note that if we follow Larsson et al.(2019), S. fuscoindicus is Hydnellum fuscoindicum, Sarcodon scabrosus is Hydnellum scabrosum, S. imbricatus and S. squamosus remain in Sarcodon, S. atroviridis may have a different status, and both S. stereosarcinon and S. calvatus are undetermined [but on spore size criteria are more likely to be in Hydnellum].
Hydnellum – The cap is tough, fibrous, with indeterminate growth – stopping then resuming with weather changes, often surrounding twigs, leaves, cones. Cap and/or stipe can be fused. K. Harrison said that it is difficult to draw a sharp line between Sardodon and Hydnellum, two genera under all conditions of growth. 8 or 10 species.
H. caeruleum – white cap margin tinted blue when actively growing; cap flesh blue-zoned, stem flesh reddish.
H. aurantiacum – at first pure white, turbinate and tomentose, changing to orange to rusty brown to brownish black, plane, corrugated and glabrous.
H. peckii – whitish with dull red cast, finally brown, exuding red droplets in wet weather, very fragrant (similar to and confused with H. diabolus).
H. suaveolens – squat, white changing to gray-brown; very pleasant fragrance while growing and on drying.
H. regium – large complex cap of numerous rosettes, deep violet black.
H. scrobiculatum – brownish caps, usually fused, with rough, deeply pitted surface; stipe base bulbous, thickened.
1a Fruitbody whitish and jelly-like, stem off-center
................................................................................Pseudohydnum gelatinosum
1b Fruitbody not whitish and jelly-like, stem central or off-center
................................................................................2
2a Fruitbody grows on wood or cones
................................................................................3
2b Fruitbody grows on soil or duff
................................................................................4
3a Fruitbody has framework of branches with spines (no true cap)
................................................................................Hericium
3b Fruitbody grows on cones (usually Douglas Fir in PNW)
................................................................................Auriscalpium vulgare
4a (2b) Spores white
................................................................................5
4b Spores brown
................................................................................7
5a Flesh tough, fibrous, thin, growth indeterminate (stopping then resuming with weather changes), caps often fused
................................................................................Phellodon
5b Flesh firm to brittle, thick, growth determinate, caps single
................................................................................6
6a Cap color pale shades of cream or orange
................................................................................Hydnum (formerly Dentinum)
6b Cap color shades of brown or gray (infrequently encountered)
................................................................................Bankera
7a (4b) Flesh firm to fragile, growth determinate, medium to large
................................................................................Sarcodon (formerly Hydnum)
(see Key to Sarcodon)
7b Flesh tough, fibrous, growth indeterminate, often fused, small to medium
................................................................................Hydnellum
(Key to Hydnellum under construction)
101a Cap not becoming scaly, growing under pine, often with pine needles adhering tightly to soft surface of cap, (not reacting to KOH when dried)
................................................................................Bankera fuligineoalba
CAP 5-13 cm, convex becoming flat to depressed or when old umbilicate, margin incurved, lobed, wavy; dull dark brown progressing through light brown to fawn and to vinaceous fawn to the margin (also described as becoming red-brown to yellow-brown), darker when wet; unpolished, when dry with adherent debris partly due to binding by surface hyphae, rarely diffracted scaly. FLESH brittle to slightly fibrous, white or tan to vinaceous buff, not zoned, in stem fibrous, pale vinaceous-buff, fawn or tan. TEETH 3-5(8) mm long, up to 0.5 mm wide, slightly decurrent, close, white or whitish becoming light red-brownish to gray-brownish. STEM 2-5 cm x 1-3 cm, colored as cap, lightest in color next to the teeth, unpolished, smooth, finely tomentose. ODOR when fresh not distinctive, when dried the odor distinctive (as in most species of Phellodon), intensely like Maggi seasoning. TASTE mild. FRUITING single to scattered or gregarious, usually under pine. CHEMICAL REACTIONS when dried not reacting to KOH and Melzer's reagent (Harrison 1968). MICROSTRUCTURES spores 4-5(6) x 4-5 um (Harrison 1968), 4.5-5.5 x 2.7-3.5 um excluding ornamentation (Breitenbach & Kränzlin), nearly round, coarsely echinulate (with spines), colorless, inamyloid; basidia 4-spored; no cystidia; no clamps; cap surface a cutis.
101b Cap bald to lacerate scaly, growing under various conifers, surface not holding debris conspicuously, (turning dark olivaceous with KOH when dried)
................................................................................Bankera violascens
CAP 3-13 cm, depressed, flat, or lobed and wavy; grayish brown or pale pinkish brown or pale purplish brown, dull brown to dark brown when old, whitish to pale fawn on margin; at first bald and unpolished, becoming finely tomentose or diffracted or lacerate scaly with scales small and appressed. FLESH soft, pallid to light brown or tinted faintly pinkish or lilac, not zoned; in stem apex as in cap, harder downward, browner to dark umber (or tinted pinkish or purplish) at base when old. TEETH 5-6 mm long, 0.5 mm wide, decurrent, fairly close, white to pallid, becoming pinkish buff or grayish. STEM 3-10 cm x 1-2 cm, single or sometimes the upper third branched, darker than cap, colored like cap but usually darker brown, often whitish near top and dark brown at the base when old; smooth, often becoming lacerate scaly. ODOR faintly fragrant, strongly fragrant when dried, (Harrison 1968), fragrant like maple syrup when fresh, soon disappearing after picking, (Bessette), pleasant, when dry intensely like Maggi seasoning, (Breitenbach & Kränzlin). TASTE mild. FRUITING single, gregarious, to cespitose under conifers. CHEMICAL REACTIONS cap and flesh instantly dark olive-green with KOH (Bessette), KOH turns dried flesh pale olivaceous, (Harrison). MICROSTRUCTURES spores 4-5.5 x 4-5 um, nearly round, coarsely echinulate, inamyloid; cystidia not seen; no clamps.
Arora (1986) discusses nomenclatural intricacies in this genus. Hericium erinaceus (lion’s mane Hericium) is easily distinguishable because it is unbranched, usually with long spines. There are three other recognizable species in the Pacific Northwest, one white to salmon-buff when fresh, with clustered spines, and growth on conifers (conifer coral Hericium), a similar one that prefers hardwoods, and has slightly longer spores, (hardwood coral Hericium), and one with spines arranged in rows along the branches like teeth on a comb, growing on hardwoods (comb Hericium). The approach of Ginns (1993) is used here, in which the comb Hericium (H. ramosum as referred to by Arora) is called H. coralloides (Scop.: Fr.) Pers. and the hardwood coral Hericium (H. coralloides as referred to by Arora) is called H. americanum Ginns. All four species have mild ODOR and TASTE. There is a negative reaction in each case when KOH is applied to the tissues, but they give an amyloid reaction to Melzer’s reagent.
201a Fruitbody unbranched, consisting of a tough piece of tissue from which spines 2-4(7) cm long hang down, growing on hardwoods
................................................................................Hericium erinaceus
201b Fruitbody branched, spines usually less than 2 cm long hanging down from branch tips or along the branches
................................................................................202
202a Spines arranged in rows along the branches like teeth on a comb, branching open rather than compact, growing on hardwoods
................................................................................Hericium coralloides
202b Spines arranged in clusters, mostly at the ends of branches, growing on conifers or hardwoods
................................................................................203
203a Growing on conifers, white to salmon-tinged or yellowish-tinged when fresh, spines up to 25 mm but usually 5-10 mm, (spores 4.5-5.5 x 4.0-4.5 um)
................................................................................Hericium abietis
203b Growing mainly on hardwoods, white when fresh, (spores 5.5-7.0 x 4.5-6.0 um)
................................................................................Hericium americanum
301a Cap yellow brown to dark brown, strongly zoned; flesh brown
................................................................................Phellodon tomentosus
301b Cap blackish, purple-black, blue-black, or dark brown, not strongly zoned; flesh blackish, purple-black, blue-black, or purple-gray
................................................................................302
302a Cap blue-black to purple-black with margin often slightly paler or purpler, flesh similar in color, spines gray to dark purple-gray-brown; relatively robust; (dark granules in Melzer’s reagent in flesh but not surface layer of cap, flesh reaction in KOH deep bluish black)
................................................................................Phellodon atratus
302b Cap usually dark brown with pale margin, flesh brown or purple-gray to purple-black; spines whitish to gray; relatively delicate; (dark granules in Melzer’s reagent in surface layers of cap but not in flesh, flesh reaction in KOH dark olivaceous)
................................................................................Phellodon melaleucus
401a Cap convex, flat or depressed, up to 17 cm across, stem 1-3 cm thick
................................................................................Hydnum repandum group
401b Cap with central depression, up to 6.5 cm across, stem usually less than 1 cm thick
................................................................................Hydnum umbilicatum group
See Note under Hydnum in the Introduction. The description here is before Niskanen et al.(2018) CAP 2-6.5 cm, convex to flat with fairly abrupt central depression (umbilicate), often with the depression continuing into the stem, surface and margin wavy; pale yellow to pale orange-yellow, becoming deep orange or reddish orange or brownish orange, slowly bruising orange; bald or slightly felted. FLESH whitish, bruising orange. TEETH up to 6 mm long, not decurrent, whitish or pale ochraceous, bruising pale orange. STEM up to 8 cm long, mostly under 1.0 cm wide, 0.45-0.7 cm wide near top, nearly central to less often off-center, may narrow slightly downward, base slightly swollen; slightly paler than cap, bruising orange. ODOR and TASTE mild. FRUITING single or gregarious, on ground in duff of coniferous forests. SPORES 9.0-10.0 x 7.0-8.6 um, nearly round, smooth, inamyloid, (Hall 1971), 7.5-9 x 6-7.5 um, nearly round, tuberculate, smooth, (Phillips), 7.5-9.5 x 6-7.5 um (Bessette). REMARKS In addition to the differences mentioned in the key lead, Hydnum repandum tends to have a lighter cap color with less red tone, more irregular outline, and more robust stature (thicker flesh and relatively shorter stem). Spores are usually smaller but there is a var. macrosporum nom. prov. with spores 9 um or longer. | Hydnum umbilicatum Boleslaw Kuznik (MykoWeb) |
GENUS AND SPECIES | KEY ENTRIES |
AURISCALPIUM Gray | |
A. vulgare Gray | 3b |
BANKERA Coker & Beers ex Pouzar | |
B. fuligineoalba (J.C. Schmidt: Fr.) Pouzar | 101a |
B. violascens (Alb. & Schwein.) Pouzar | 101b |
= Bankera carnosa (Banker) Snell, E.A. Dick, & Taussig | |
HERICIUM Pers. | |
H. abietis (Weir ex Hubert) K.A. Harrison | 203a |
H. americanum Ginns | 203b |
H. coralloides (Scop.: Fr.) Pers. | 202a |
H. erinaceus (Bull. ex Fr.) Pers. | 201a |
HYDNELLUM P. Karst. (Key to Hydnellum under construction) | |
H. aurantiacum (Batsch: Fr.) P. Karst. | Introduction |
H. caeruleum (Hornem. ex Pers.) P. Karst. | Introduction |
H. peckii Banker | Introduction |
H. regium K.A. Harrison | Introduction |
H. scrobiculatum (Fr.) P. Karst. | Introduction |
H. suaveolens (Scop.: Fr.) P. Karst. | Introduction |
HYDNUM L. | |
H. repandum Fr. | 401a |
= Dentinum repandum (Fr.) Gray | |
= Hydnum washingtonianum Ellis & Everh. | |
H. umbilicatum Peck | 401b |
= Dentinum umbilicatum (Peck) Pouzar | |
PHELLODON P. Karst. | |
P. atratus K.A. Harrison | 302a |
P. melaleucus (Fr.) P. Karst. | 302b |
P. tomentosus (Fr.) Banker | 301a |
PSEUDOHYDNUM P. Karst. | |
P. gelatinosum (Scop.: Fr.) P. Karst. | 1a |
SARCODON Quel. ex P. Karst. (see also Key to Sarcodon) | |
S. calvatus (K.A. Harrison) K.A. Harrison | Introduction |
S. fumosus Banker | Introduction |
S. fuscoindicus (K.A. Harrison) Maas Geest. | Introduction |
S. imbricatus (L.: Fr.) P. Karst. | Introduction |
S. scabrosus (Fr.) Quel. | Introduction |
S. stereosarcinon Wehm. | Introduction |
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