A Real American Ivory-billed Woodpecker

by Leon Shernoff

Echinodontium ballouii, in mid-winter. Photo by Bill Neill

Echinodontium ballouii, in mid-winter. Photo by Bill Neill


Almost a hundred years ago, William Hosea Ballou discovered a small, strange fungus growing on an endangered tree, Atlantic white cedar, in New Jersey. With typical hyperbole, he commented “There is no fungus more beautiful – or more deadly!” However, the fungus turned out to be in considerably more danger than its host, and Ballou’s collections of 1909 were the last time anyone has seen it in the wild. Until now…

The Explorers

Larry Millman and Bill Neill are connoisseurs of obscure fungi. Bill’s perseverance in exploring and documenting the fungi of coastal Massachusetts led to his co-authorship of Mushrooms of Cape Cod and the National Seashore with Arleen & Alan Bessette. Likewise, Larry is a member of the Explorers Club, with a life-long interest in the Arctic that has led to numerous books. Each of us has our own personal reaction upon spotting a mushroom in the forest. For many of us, this is “Is it edible?” For Bill and Larry, it’s “Have I seen that before? Do I know what it is?” So their interest was piqued when they each, independently, came across a reference to a mushroom that no one had seen for almost a hundred years. Only three specimens – all collected by the same person, at the same location – have ever found their way into scientific collections: two are at the New York Botanical Garden, and one is at the USDA herbarium at Beltsville, Maryland.

The type specimen of E. ballouii, collected in 1908, at the New York Botanical Gardens

The type specimen of E. ballouii, collected in 1908, at the New York Botanical Gardens

Both men credit “the bible,” Robert Gilbertson and Leif Ryvarden’s 1986-7 Polypores of North America,  with starting them on the trail. Most of us know polypores as “those bracket fungi” that stick sideways out of trees like little bookshelves. Many of them are woody, and persist in the woods for several months – some are even perennial, with a new layer of spore-bearing tubes coming out repeatedly beneath the last one. Because of this, they are well-suited for wintertime mushroom-hunting, when almost nothing else is out. Bill notes that this also fits in well with Larry’s arctic bent, and that Larry has gotten the Boston mushroom club to extend its schedule of meetings past the time of year when it holds walks, because Larry is still out finding things that he wants to share with the club. Both read in “the bible” that this polypore was presumed extinct, and as Larry noted, “I’m an explorer, so seeing something like that is like showing a red flag to a bull.”

The Fungus

The fungus they were looking for is an Echinodontium, thought by some people to be a “living fossil” genus of mushrooms. Only about half a dozen species exist worldwide. The thick, woody spines that project from the underside of the cap have long been thought to represent an early stage in fungal evolution. The past couple decades of mycological research has taught us that in fact we can’t rely on this sort of feature to tell us which fungi are more “primitive” and which are more “advanced,” and we’ll look at what this research has to tell us later in the article. But for the moment, let’s take time out to look at what an attractive narrative of evolution this older approach provides.

E. ballouii in situ. Photo by Bill Neill

E. ballouii in situ. Note how large the (very slow-growing) tree is, and how thick and blunt the fungus’s spines are. Photo by Bill Neill

Many fungi increase the surface area of their spore-bearing surface by roughening it, allowing them to produce more spores in a compact space, just as crumpling a piece of paper compresses the same amount of surface area into much less space than the same piece of paper laid out flat.

This <i>Hericium</i> has long been thought to be a more modern, "highly-evolved" tooth fungus. Its spines are long and thin, maximizing spore-bearing surface area, and its branches are also thin not "wasting" as much resources growing flesh that isn't directly bearing spores.

This Hericium has long been thought to be a more modern, “highly-evolved” tooth fungus. Its spines are long and thin, maximizing spore-bearing surface area, and its branches are also thin not “wasting” as much resources growing flesh that isn’t directly bearing spores. Photo by Leon Shernoff

More “modern-looking” fungi are very efficient at this – for example, if you took each gill of a button mushroom from the store and opened them out flat, they would cover an area the size of a trash can lid. Larry and Bill’s find, Echinodontium ballouii, seems to have only just started this roughening process – its lower surface it is merely wavy, with a few nipple-like bumps and what seem to be incipient stalactites. It very much looks like the roof of a cave. Its nearest relative, which looks very similar, is Echinodontium ryvardenii, which occurs only on hundred-year-old Juniper in Sardinia (Annarosa Bernicchia & Andrea Piga. “A new species of Echinodontium from Italy,” Mycotaxon 68: 483-491. 1998)

<i>Echinodontium ryvardenii</i>, on old-growth juniper in Sardinia. Photo by Cristina Spinelli.

Echinodontium ryvardenii, on old-growth juniper in Sardinia. Photo by Cristina Spinelli.

E. tinctorium, North Amreica's most common species of Echinodontium. Photo by John Denk. It looks almost like driftwood

E. tinctorium, North Amreica’s most common species of Echinodontium. Photo by John Denk. It looks almost like driftwood, but when you look at the interior…

The only other Echinodontium species in North America is Echinodontium tinctorium, which is limited to old-growth forests in the Pacific Northwest. It has one of the more modern-looking spore-bearing surfaces in the group, as its underside is thickly clustered with spore-bearing spines, although they are thick and blunt, and thus provide less area than the thinner ones that are more common today. E. tinctorium is also the best-known species of Echinodontium: it is prolific enough to be considered a “serious pest” of timber in the area, and its woody red and orange interior flesh was ground up and used for body paint and other ritual purposes by the Native Americans of the region.

The interior of E. tinctorium shows the flesh that made it useful as a face and body paint for nNative Americans in the Pacific Northwest. Photo by John Denk.

The interior of E. tinctorium shows the flesh that made it useful as a face and body paint for Native Americans in the Pacific Northwest. Photo by John Denk.

As we’ll see, the more recent research has produced a more complicated picture of the evolution of fungi, and we lose the nice tidy sequence presented in these pictures. But the new view of fungal history has its own surprises to unveil, and I think that it is richer and more interesting. But all this is later on. While Bill and Larry were on their quest, they still thought they were looking for a dinosaur of the fungal world.

The Search

Bill and Larry discovered their mutual interest on a walk a few years ago, when they found themselves at the Ponkapoag Bog near Boston. Ponkapoag is one of the most accessible bogs containing a tree commonly called Atlantic white cedar (although it is not closely related to true cedars), Chamaecyparis thyoides. Echinodontium ballouii occurs only on this one tree – itself a “living fossil.” Atlantic white cedar was on the verge of extinction a hundred years ago: highly valued for ship-building because of its attractive white wood and resistance to rot, it was rapidly being cut down (as it had for centuries). Even in that environmentally insensitive time, its extinction was considered likely within a decade. With the increasing switch to metal-hulled boats, however, it fell by the wayside as an industrial product, and recent efforts to re-establish it along the Atlantic coast have been very successful. Ponkapoag is one of the spots where it is now growing, and the pair of myco-explorers decided to take a shot at finding its associated polypore there. They quickly realized, however, that the site was unsuitable, and gave it up. “It had clearly been logged quite thoroughly,” said Bill. “It was all young trees, and we thought that the mushroom was going to need old growth.”

And thus commenced the search. How to find the proper cedar swamp? “Bill did a lot of great work on Google Earth, finding cedar swamps and looking at the canopy, getting a feel for whether this might be the sort of place that would be worthwhile to try,” says Larry. Then it was off in the middle of winter to explore the candidate locations. Why the middle of winter? Well, winter in New England was a natural for an Arctic explorer, of course. But it also made it possible to get to places in the cedar swamps that were otherwise inaccessible.

“Snowshoes are not a common mycological tool,” notes Larry. “But here they were indispensable. In summer or fall these places are mucky, wet, hot, bug-ridden and impenetrable. You have to go in the dead of winter when the water’s frozen in the swamps. The snowshoes were our big breakthrough.

“The first time we found it was a cold, snowy day,” says Larry, “with what New Englanders call a flirt of snow. We had visited one or two sites earlier on, and now the snow was starting to accumulate, so we were thinking about going back. But we said what the hell, let’s go for it.”

Echinodontium ballouii, showing its stubby spines and often tenuous attachment, along the underside of a branch stub.

Echinodontium ballouii, showing its stubby spines and often tenuous attachment to the tree, along the underside of a branch stub. Photo by Bill Neill

“It was really grey, with some deep snow by then,” Bill confirmed, “and not having found anything at the first place, we didn’t have a lot of enthusiasm any more. But when we walked in, we started to see some fungi, which was unusual – in the other cedar swamps there weren’t many, but here there were some even on the cedar trees. We walked in circles, and for at least the first hour we didn’t see anything. But we were pretty relentless by then – we had driven all this distance, and decided to look until it got dark. We were focusing on branch stubs as a possible site for it. That’s not listed in North American Polypores, but Echinodontium tinctorium does that, using the branch stubs as entry wounds.”

“It was eerie, with a mist, and a raven hovering above. I regard ravens as an icon of good luck, a sort of totem bird for me,” says Larry. “We reached some trees that were larger than the ones in the other swamps, and this was a good sign.”

“At one point Bill saw a blackened mass at about eye level on an old gnarled tree. He wondered if that was it, and I was skeptical. Bill didn’t know what else it could be – there was no other possibility. We each walked around to other side in our own direction, and Bill found the first specimen that was obviously E. ballouii, growing on other side. It was a celebratory moment. There truly was nothing else it could be. We continued to look as it was getting dark, and found another half dozen. We decided to come back, but not to collect them, and GPSed each tree. Unfortunately, as I was taking a photo of one, my camera bumped it and it fell off – Ballou mentions this: that they’re very precariously attached to the tree. Bill took it to the New York Botanical Garden to get its identity verified. We were both tremendously excited.”

“I said to Larry, ‘You’re not going to believe this.’ I had never seen it before – neither of us had – I just had the description, the concept in my mind. It doesn’t look much like E. tinctorium – it has these encrusted teeth that look like dripping wax or stalactites. It’s not a fine-toothed entity like E. tinctorium or other ‘tooth fungi’.”

“The ‘teeth’ on the underside are just bumps, distinctly nublike,” says Larry. “Ballou said that his were golden yellow underneath and then faded, but ours are sort of drab pink.  The top of it is almost a camouflage coating of mosses and lichens. It’s a shaggy mushroom on a shaggy tree.” Bill agrees that “The bark of all these trees is just a wonderland of lichens and similar organisms. The Echinodontium fits right in.” He also suggested that the difference in hymenium color might be because Ballou’s specimens were collected during the growing season.

Another unusual feature of the fungus is that it often occurs quite high up in the tree. “‘Echinodontium neck’ is definitely an occupational hazard when looking for these things,” says Larry. “Ballou talks about his specimens coming from 40-50 feet aloft,” Bill confirms. “When he found his, he was climbing trees looking into bird nests. Then he immediately goes on to state that this is the typical way these things grow, from just the few collections that he made. But of course the first ones we found were at eye level.”

Sometimes the fungus sticks a cap out from the tree; at other times, the spore-bearing surface is plastered directly on the bark. It did indeed show a preference for coming out just below a branch stub, just as Bill and Larry expected, but while it may indeed use the exposed wood as a way of getting into the tree, that doesn’t mean that it has to fruit there – after all, the mycelium can extend for quite a ways. I think that it’s more likely that the fungus just isn’t very good at forming a cap on its own, so it’s easier for it to spread out horizontally be extending along the bark on the underside of a branch stub, forming a cap that way and just sticking out a little from the branch on each side.

Maggie Rogers weighed in with some confirming data from the West coast, when she heard that the rediscovered mushroom was from the same genus as the Indian Paint fungus, E. tinctorium: “Dunno if you’ve ever seen Echinodontium in situ: they weigh a lot! You don’t want to be beneath when they let go. I’ve been collecting them for the January meeting of the International Fiber and Fungi Institute meeting in Mendocino, since they’re not found often elsewhere. We get rather huge specimens… if one fell on your head, you wouldn’t live through the experience. The largest I’ve held weighed upward of 10-15 pounds. They, like their cousins, tend to hang higher up in the trees, though if a major branch stub is lower, there’s a chance for a lower infection.”

My dad pointed out that regardless of how closely E. tinctorium and E. ballouii turn out to be related, a similar physical problem (precarious attachment to the tree) could have resulted in a similar solution (growth under branch stubs). Having the top of the cap attached to the lower surface of a branch stub helps it hold on for as long as possible! Hopefully, E. ballouii will become well enough established that we can take a more thorough look at it and figure out its evolutionary relations.

The Ecology

It’s impossible to know how many trees are now inhabited by Echinodontium ballouii, because the main body of the fungus would be hidden deep within the tree. This mycelium, which resembles a network of fine white roots, weaves its way through the wood, dissolving and digesting it as it goes. The spiny fruiting bodies that we see outside of the tree are only sent out when the fungus wants to reproduce, dropping its spores on the wind to land on another Atlantic white cedar and start another Echinodontium mycelium.

The west coast species of Echinodontium is thought to spend an unusually long time inside the tree (around seventy years) before sending out their fruiting bodies, which makes them especially vulnerable to human activities that kill the trees while before the conk emerges. Other research from the Pacific Northwest has led some foresters there to believe that the fungi that appear on the old-growth trees there can live inside them for upwards of 300 years, forming conk after conk, before the tree comes down.

“My feeling for the Atlantic white cedar is that the tree has to be over a hundred years old for the fungus to appear,” says Larry. “Perhaps not real ‘old-growth,’ but certainly quite old. When the tree is chopped down before it reaches a certain size, this prevents the fungus that has co-evolved with it from ever appearing on the outside, thus cutting off the next generation of fungi. The same goes for hydrology changes that kill the trees at a young age, and of course just cutting them down to make room for shopping malls.”

Dead crown of an Atlantic white cedar tree, with its branches thickened by cankers caused by Gymnosporangium biseptatum. Photo by Bill Neill.

Dead crown of an Atlantic white cedar tree, with its branches thickened by cankers caused by Gymnosporangium biseptatum. Photo by Bill Neill.

Both Bill and Larry started out by trying to use the 1988 book Atlantic White Cedar Swamps in New England as a guide to locating the trees; but in spite of its fairly recent date, the book proved to be almost useless. “Since then, in many locations the water has risen and drowned the trees,” says Larry. “Or they have dried up completely. One site in western Massachusetts was supposed to be a classic Atlantic white cedar swamp, but now it’s mostly spruce and sugar maple. We took one look and didn’t even bother going in. And just last month I was up looking in Appleton, Maine. There were a lot of old moss-covered stumps, lots of young growing alders and birch, and no E. ballouii in sight.”

“At one time, it was probably in every cedar swamp,” notes Bill. “Some of these have been well-preserved and are even interesting historically: for instance, one on Cape Cod, is actually the site where Marconi made his first radio transmission. It’s been beautifully kept and the trees have grown back – some of them are probably two hundred years old. There’s even a boardwalk where you can walk in through what would otherwise be really rough territory and see them. But most of these sites have been totally raped.

“New Jersey used to have huge areas of cedar swamp. I read one account from the 1600s: someone from Europe visited where the Meadowlands are now. It was filled with gorgeous giant cedars. The visitor said that they shouldn’t be cutting them all down, because they were so beautiful. But now of course they’re called the Meadowlands because there are no trees left of any kind.

“New Jersey had a major industry back then in what was called ‘bog iron.’ The acidic, peaty water dissolves iron from underground deposits, and if you boil the water off you can precipitate the flocculent iron. This was a very important source of iron for farm tools, guns – they even made cannons for the Revolutionary War from bog iron. They just cut down as many of the trees as they could right there to burn to boil the water, and huge other areas were razed to make charcoal for the bog iron industry. Besides that, it was great for shingles and boats, because the wood was so weather and rot resistant, and had this attractive white color. It was tremendously popular. One piece of literature mentions a spot where the trees had been totally cut down to ground, and speculated that it might be economical to mine the stumps that were pickled in the earth.

“So how can a fungus that doesn’t fruit except from old growth trees continue to survive? We only found it in a few places in one swamp, out of all the locations that we went to – these isolated pockets of just the right sort of situation. And who knows what vector has spread it, historically? Perhaps there was a woodpecker, or some other sort of bird, that was primarily responsible for carrying the spores from one tree to another, and with the swamps so far apart, that’s not going to be possible anymore.”


The co-existence of Echinodontium and Chamaecyparis species may provide material for some interesting ecological studies. Gross included six species in his delineation of the genus Echinodontium in 1964. There are currently five species of Chamaecyparis in the world. They grow in exactly the same places, and in two of those places the Chamaecyparis tree is the only host of the local Echinodontium species.

Two of Gross’ Echinodontium species have been transferred in the genus Laurilia (where they are the only two species), because they look somewhat different to the eye. Gilbertson and Ryvarden point out that their great similarities in microscopic morphology support Gross’s practice of retaining them in Echinodontium, and this host association provides another layer of support. Let’s take a closer look at the world-wide locations of Echinodontium and Chamaecyparis.

North America’s Pacific Northwest is home to Chamaecyparis lawsoniana, ranked eleventh-largest tree in what is truly a stupendous habitat for large trees. It is also home to Echinodontium tinctorium. North America’s Atlantic coast, as you know, is home to C. thyoides and E. ballouii. Japan is home to two species of Chamaecyparis and two of Echinodontium. And in Taiwan, Chamaecyparis formosensis is the sole host for Laurilia taxodii. So it looks a bit like Echinodontium may have started out exclusively on Chamaecyparis, and eventually jumped to other nearby trees in Japan and the Pacific Northwest.

Two fungal species are the odd men out: Laurilia sulcata is found across the boreal zone, wherever there are giant spruces (in Europe, exclusively on Norway spruce, Picea abies); and the newly discovered E. ryvardenii is found on Sardinia, growing on juniper a hundred years old and older. So we know that Echinodontium in Gross’s sense (including Laurilia) likes ancient conifers, with clearly a vastly disproportionate affection for Chamaecyparis. But what does the more modern evidence say? That gets more complicated, and we’ll look at that separately in the article on DNA.

The Future

Unfortunately, one more similarity between the various locations of Echinodontium and Chamaecyparis is the over-exploitation of Chamaecyparis trees for timber. The uses of Chamaecyparis thyoides have already been covered. In east Asia, where buildings are lower and humidity is high, the trees were immensely valuable for use in building temples and palaces – wherever gorgeous, long-lasting wood was a priority. Now that the Japanese and Taiwanese stock of these trees are almost exhausted, demand for this sort of wood has made the Pacific Northwest’s C. lawsoniana the most valuable wood in the region. Almost all of it goes for export to east Asia. And if this weren’t enough of a problem for the existing North American stock, we’ve managed to introduce a foreign Phytophthora (yes, just like Sudden Oak Death), P. lateralis, that is killing all the C. lawsoniana in the wet areas that it likes most. The USDA forestry service’s web page discussing the tree is a bit obtuse about what is the most pressing issue here:

“This valuable tree, however, has a very limited range and an uncertain future. Management of Port-Orford-cedar [C. lawsoniana] has become impossible in much of its range since the introduction of a fatal root rot that is still spreading. Old-growth forests are being depleted rapidly, and the use of second-growth forests is complicated because early growth is relatively slow. The commercial future of one of the most beautiful and potentially useful trees will depend on development of silvicultural practices that minimize infection by root rot.”

Apparently simply not cutting down the trees has not yet occurred to the feds. And in fact, we see from their page on C. thyoides that – now that the tree itself seems to be out of ecological danger – it is again listed as just another commercial wood-bearing tree, even in the same breath as noting its imminent destruction a hundred years ago:

“Heavy cutting for many commercial uses during this century has considerably reduced even the largest stands so that the total volume of this species growing stock is not currently known. It is still considered a commercially important single species in the major supply areas of North and South Carolina, Virginia, and Florida.”

You can even get a rundown of the various attractive features of its wood on another US Forest Service page:

The sapwood of Atlantic white cedar is narrow and white, while the heartwood is light brown with a reddish or pinkish tinge. The wood has a characteristic aromatic odor when freshly cut and has a faint bitter taste. It is light weight and has a fine texture and a straight grain. It is moderately soft, low in shock resistance and is weak in bending and endwise compression. It is very resistant to decay, works easily with tools, shrinks little, finishes smoothly, holds paint well and splits easily.

Perhaps one more interesting fact about these trees will emphasize the importance of treating them as parts of an ecosystem and not as reservoirs of standing board-feet: you know these trees. I know that it seems from this article that they’re these exotic, rare organisms, but in fact if you’ve seen any professionally planted shrubbery at all, I can almost guarantee that you’re familiar with them – it has proved very easy to produce cultivars of different shapes and sizes, and their extremely slow growth means that they stay “as they were put” for much longer than a regular tree that’s going to shoot up and overtake your house. And of course, the tremendous resistance to fungal decay is also a plus. Chamaecyparis trees’ needles form flattened fronds of tiny, overlapping scales, and produce tiny purplish, reddish-brown or pale grayish-blue cones that are so small that they look like berries. If you can’t immediately recollect them in your neighborhood, keep an eye out and it won’t take you long to spot them.

So the trees in the abstract are in no danger of going extinct – C. lawsoniana, which has been so suddenly threatened by this introduced Phytophthora, is one of the most popular yard shrubs around. However, this dramatizes the difference between these trees as ornamentals and in their native ecosystems – Christopher Earle’s conifers.org website lists the largest known existing C. lawsoniana as being 229 feet high, with three taller ones over 70 meters high, and he mentions stumps from before the era of recording these statistics as being six meters (just short of twenty feet) across. He gives the standard mature dimensions for C. lawsoniana at present as 50 meters tall and 3 meters diameter at chest height. Needless to say, at that size it’s also covered with other organisms (and their associated organisms) and is a major hub, as all giant trees are, in its ecosystem.

It’s not just Echinodontiums that only live on these trees – C. thyoides is also the sole food source for the caterpillars of the rare Hessel’s Hairstreak butterfly, which only occurs in the Atlantic white cedar swamps because the flowers that the adult butterflies like only grow there. And, quite frankly, there are probably hundreds of other similarly unique organisms associated with these unusual trees. Of the fungi that are large enough to see, across the country, knowledgeable estimates are that we’ve named only 5% to 20% of what’s out there. That figure goes way down for inaccessible areas like Atlantic white cedar swamps! And, especially, research in the coastal redwood forests and the Great Smokies have taught us that all sorts of communities and mini-ecosystems exist in the canopies of these dense, damp forests. But today, even in the Atlantic white cedar swamps that are still standing, there’s little of the old canopy left. Who knows how many of the original canopy organisms are still alive?

There is more hope for fungi than for most other associated organisms, because of the fungal life-style. “We don’t know how many trees have the mycelium growing in them,” Larry points out. “If we came back in a hundred years, would we see lots of E. ballouii? More even than the conks that we now regard as common? Scott Redhead rediscovered Mycena gaultheri in the 1980s, fifty years after it was listed as extinct. So all sorts of things are possible. But we can kill these things so easily by destroying their habitat that we shouldn’t be too optimistic.”

Greg Mueller says “For me, that’s the great thing about this discovery: it highlights how many unique organisms are still at large in our old-growth forests, and how important it is to get out there and look for them, and to preserve these habitats. Sabine [Huhndorf, another prominent mycologist at the Field Museum] went out to that one pocket of old-growth forest remaining on the upper peninsula of Michigan last year, and she also found several things that no one has found for seventy or eighty years. It’s always great to see something rediscovered like this, and it emphasizes that we still need to explore and preserve the few areas of eastern old-growth forest that are left. Mycologists from even fifty years ago didn’t bother to designate their finds as being old-growth specific, because that wasn’t even an issue in their minds; but it’s turning out to be quite important for some species, and is undoubtedly equally important for other organisms besides fungi.”

And Roy Halling, the chief mycologist of the New York Botanical Garden, says “To my mind, this is one of the most significant events for field mycology in quite some time. Such a find indicates again that basic field work is the foundation of all of biology.”

In honor of Bill and Larry’s discovery, we are dedicating next issue’s installation of “America’s Most Wanted Fungi” to other polypores that are listed by Gilbertson & Ryvarden as only having been found once or a few times. One such fungus – Abortiporus fractipes – has already been featured, with the news that it’s not as rare as once thought – it’s just that people haven’t been looking for it. We hope that this article will inspire Mushroom the Journal’s readers to go out and find other things that nobody else has seen in a long, long time.

Related content

Echinodontium ballouii: from eyeballs to DNA
William Hosea Ballou (biographical sketch)

This article first appeared in the Fall 2007 issue of Mushroom the Journal