Mushroom Identifier

Dark Honey Fungus

Armillaria ostoyae

Also called armillaria solidipes, shoestring root rot, dark armillaria

A robust clustered honey fungus of conifer forests with a dark-scaly reddish-brown cap and a thick white ring edged with brown scales, spreading through the soil by black shoestring rhizomorphs.

Overview

The Dark Honey Fungus is the conifer-associated member of the Armillaria complex and the most robust and darkly pigmented of the group. It fruits in large dense clusters at the bases of standing trunks, on stumps and directly from buried roots, sometimes producing scores of caps in a single tight tuft.

Armillaria is a genus of white-spored, ringed, wood-inhabiting fungi that were long lumped together as a single variable "honey fungus" until careful work on mating compatibility and morphology separated them into a series of biological species. A. ostoyae — also published under the name A. solidipes — is distinguished by its dark scales, its thick ring with a brown-scaled edge and its strong association with conifers.

The species is famous for producing some of the largest individual organisms ever documented: a single genetic individual in the Malheur National Forest of eastern Oregon covers several square kilometres of forest soil, all connected by a network of black cord-like rhizomorphs radiating from the parent mycelium.

How to identify it

Cap

The cap is 4–15 cm across, convex to bell-shaped when young, expanding to broadly convex or flat with a wavy margin. The ground colour is a warm pinkish-brown, reddish-brown or dark tan, and the surface is covered — especially over the centre — with prominent, erect to recurved dark brown or blackish scales that thin out towards the rim. The scales are dry and persistent, giving the cap a rough, bristly texture quite unlike the smooth-capped members of the genus.

Gills

The gills are adnate to slightly decurrent, moderately crowded, and whitish to cream when young. With age they discolour to dull pinkish-tan and often develop rusty or dark brown spotting, particularly along the edges and where bruised.

Stipe

The stipe is 5–15 cm long and 1–2.5 cm thick, firm and fibrous, whitish above and darkening to brown below, often tapering or fused with neighbouring stipes at the base. It carries a thick, persistent, cottony white ring high on the stipe, and the diagnostic feature is the ring's underside and edge, which is decorated with brownish to blackish woolly scales or a distinct dark band. Below the ring the stipe frequently shows further bands of brownish scales.

Flesh and spore print

The flesh is firm and white, sometimes browning slightly where cut, with a somewhat astringent smell. The spore print is white — an important separator from the many brown-spored fungi that fruit on the same wood. Black, flattened, bootlace-like rhizomorphs can usually be found under the bark of colonised roots or trunk bases.

Where and when it grows

The Dark Honey Fungus is overwhelmingly a conifer species, associated with spruce, fir, Douglas fir, pine, hemlock and larch, and only occasionally found on hardwoods. It fruits in dense caespitose clusters at the base of living and dead trunks, on stumps and from the soil directly above infected roots, so clusters may appear a metre or more from the nearest visible trunk.

It is widespread across the boreal and montane conifer forests of North America, Europe and Asia. Fruiting is concentrated in late summer and autumn, generally beginning after the first substantial autumn rains and continuing until hard frosts, and in a good year a single infected stand can carry clusters at the foot of many trees at once.

Ecological role

This is a facultative parasite as well as a wood decayer. Its rhizomorphs — tough, black, cord-like structures resembling bootlaces — grow through the soil in search of new roots. On contact, the rhizomorph can penetrate the root bark and establish in the cambium, producing sheets of white mycelium between bark and wood that spread up into the root collar and lower trunk. Once the tree dies, the fungus continues as a saprotroph, causing a white rot of the roots and butt.

Because the mycelium can persist in stumps and root systems for decades and expand outwards year after year, individual clones reach extraordinary sizes and ages, with the Oregon A. ostoyae individual estimated at thousands of years old. Ecologically the species is a major agent of natural thinning in conifer forests, creating canopy gaps, standing dead trunks and coarse woody debris that many other organisms depend on.

Similar species

Armillaria mellea has a honey-yellow to olive-brown cap with only sparse fine fibrillose scales concentrated at the centre, a thick white ring without a dark scaly edge, and fruits in very dense clusters chiefly on hardwoods. Armillaria gallica has a paler tan cap with fine yellowish scales, a markedly bulbous stipe base and a fragile, cobwebby yellow-fringed ring that often disappears entirely.

Pholiota squarrosa is superficially similar with a dry scaly cap and clustered growth at tree bases, but its cap is dry ochre-yellow with greenish-tinged gills and it gives a rusty-brown spore print rather than white. Galerina marginata on the same conifer wood is far smaller, smooth-capped, with a thin ring and rusty spores.

Common questions about Dark Honey Fungus

What are rhizomorphs and where do I look for them?
They are dense cords of mycelium with a tough black rind, resembling bootlaces or shoelaces. Peel back loose bark at the base of an infected conifer, or dig gently at a colonised stump, and you will often find flattened black cords running vertically beneath the bark, along with white sheets of mycelium in the cambium.
How do I tell it from Armillaria mellea in the field?
Look at the cap scales and the ring. *A. ostoyae* has coarse, dark brown to blackish erect scales on a reddish-brown cap and a thick white ring whose edge carries brown to black woolly scales. *A. mellea* is honey-yellow with much finer, sparser scales and a plain white ring with no dark banding, and it favours hardwoods.
Is it really the largest organism on Earth?
One clone of this species in the Malheur National Forest in Oregon covers several square kilometres of forest floor and is one of the largest single organisms ever measured by area. The whole extent is a single genetic individual connected by rhizomorphs, though the visible fruiting bodies are scattered clusters.
Why do clusters sometimes appear away from any tree?
The fungus fruits from the roots it has colonised, not only from the trunk base. Because conifer root systems extend well beyond the crown, and because rhizomorphs travel through soil, a cluster can push up through litter several metres from the nearest standing trunk or from a stump long since removed.