Sunday, 23 October 2016

The Piggyback Pinkgill: The Mushroom Mushroom-Parasite Volvariella surrecta

Last week's Mycological Society of Toronto foray was a wet one. It rained for hours. Despite the amount of water that was falling from the sky, there were slim pickings since it's been unseasonably and unreasonably dry for months. 

Because of this dearth of fungi, I was doing a quick last minute gleaning of the woods beside the parking lot hoping to add to the meagre display on the identification table when—through my dripping glasses—I spotted a tight clump of whitish mushrooms among the fallen leaves. My first thought was that they were Coprinoid. Knowing that I wouldn't be bringing it home if they weren't easily identified (I knew they'd liquify before I'd have time to get to them), I just roughly grabbed a couple as a sample. 
The first engraving of Volvariella surrecta by John Leonard
Knapp, 1829, showing strangely unaffected hosts.
The host of the ones I found, Leucopaxillus albissimus,
was barely recognizable as a mushroom.
Since the mushrooms came up attached to a squidgy piece another, larger mushroom, I knew before I even straightened up what I'd found: Volvariella surrecta (though its name escaped me and I had to be reminded of it by Nathaniel, our club's 13-year-old budding myco-meister!). Nonetheless, it was a joyful moment—but a sad one, too. Joyful because I've been wanting to find this character for nearly a quarter century. But sad because the specimens were pristine, with multiple stages of development—which would have been the perfect set-up for photography of a rare species. If I hadn't thoughtlessly wrecked its display. Dang!

Volvariella surrecta emerges from a well-developed volva, or cup.
Volvariella surrecta is one of very few mushrooms that grow on other mushrooms. It's pretty picky about who it grows on—primarily on Clitocybe species, but occasionally Tricholoma. Apparently in western North America and in Europe. Its preferred host is Clitocybe nebularis, which is not found in eastern North America. 

Volvariella surrecta is usually found on Clitocybe nebularis. (Wikipedia)
There's some dispute about whether it's an obligate parasite or if it's sometimes simply saprobic on its hosts. Some host fungi are considerably deformed, but are otherwise solid, while others, like mine, are decaying messes. I should say that I had a devil of a time finding any spores on the seemingly undistorted gills of the Leucopaxillus albissimus that mine had chosen to grow on. I eventually found a dozen are so. I have no idea if they would have been viable or not. 

Though rare, Volvariella surrecta is easy to recognize.
There were lots of little nubbin babies waiting in the wings.

On-line References for Volvariella surrecta:

on Mycoquebec
on Scottish Fungi (are the Scots responsible for the common name "Piggyback Pinkgill")
Arthur Henry Reginald Buller (1924) Researches on Fungi: vol. 3

Wednesday, 17 August 2016

Nature's Spray Paint: Inonotus glomeratus

fallen tree covered in yellow spores

I've never been able to pick favourites—be it a colour, a food, a flower, or a fungus. But I do have long lists of top contenders.

I have a particular soft spot for fungi that do something, that, in particular, show me process, whether it be life cycle or interaction with other Kingdoms or longterm movement through a landscape. Which is why it's a no-brainer for Inonotus glomeratus to make my fungi long list.

Here are the basics about Inonotus glomeratus:

  • it's a polypore
  • it can be found on a wide range of hardwoods 
  • its fruiting bodies show either effused-reflexed or resupinate growth
  • its spores are bright golden yellow 
  • it is both parasite and saprophyte 
But what does it do  

1. It Produces a Cool Optical Effect

Inonotus glomeratus is one of a many polypores that produces a "glancing" effect when the its pore surface is turned towards and away from light. When it catches the light the right way, it seems to flash silver.

Inonotus glomeratus glancing pore surface
This chunk of I. glomeratus shot from two different angles shows the
glancing effect on the pore surface. It's wickedly hard to photograph.

2. It Weeps Black Teardrops

tar-like black drops Inonotus glomeratus
Black, tar-like guttation droplets 
Well, they're not black teardrops; they just look like black teardrops...or blood...or tar. 

When the fungus is young and actively growing, it produces black drops of exudiate, or guttation (see my post on this phenomenon). Since I. glomeratus is usually resupinate and white to very pale grey at this stage, the black drops can be quite startling. The exudiate is often so plentiful that the ground beneath is splattered with pools of it. 

exudiate black drops polypore Inonotus glmeratus
The contrast between the lightness of the pore surface
and the black guttation can be striking.

black weeping Inonotus glomeratus polypore
Guttation droplets can be so plentiful that they drip to the forest floor.

guttation pits on Inonotus glomeratus
The droplets are sometimes reabsorbed into the fungus,
leaving negative drop-like pits.

3. It Paints Its Surroundings Yellow

polypore coats log with yellow spores
It can produce copious amounts of yellow spores.
Not only does I. glomeratus produce ridiculous amounts of spores—on par with Ganoderma applanatum—its spores are an unusually bright yellow or sulphur. When there is little drought or wind near the time of maturation, this unassuming fungus can cover everything near it with with a yellow coating that looks exactly as if someone has gone overboard with a can of spray paint. It's a particularly eerie effect when it happens in woods you know no one else goes in.

yellow spores of Inonotus glomeratus polypore
white resupinate polypore Inonotus glomeratus
These two shots were taken just a few days apart—the second, after a rain.

4. It Grows Whiskers

setae hairs pore surface of Inonotus glomeratus
Close-up of Inonotus glomeratus pores showing setae, or hairs

Like most Inonotus species, I. glomeratus grows setae, which stick out of the pore surface like minute whiskers that are visible under magnification. Under the microscope these abundant setae look more like weapons than hairs.

setae hairs Inonotus glomeratus
Under the microscope, the hairs look vicious.

5. It Can Mimic Chaga

Inonotus glomeratus canker conk looks like chaga
An I. glomeratus canker can look a lot like chaga. (Dianna Smith)

Another thing this extraordinary fungus does is produce sterile cankers that can look exactly like the infamous Inonotus obliquus canker, more commonly known as chaga. This happens in living trees, in its parasitic stage, after entering a living tree through a wound. Once it's gained entry, it causes a white rot of the heartwood. 

Inonotus glomeratus canker conk on maple
This knot is the likely entry point of I. glomeratus on this red maple.
Though I. glomeratus is most commonly found on maples and oaks, and is not uncommon on Populus, and can also be found on other hardwoods, including, I’ve read, on Betula, or birches.When it attacks maples, it does not normally develop a chaga-mimicking canker, rather it causes a prominent ridge of bark to grow around a rotten knot. Once this appears on a maple tree, (or a lumpy canker grows on another hardwood), there can already be a column of decay inside the tree 3 to 4 meters high. The fruiting bodies that produce the yellow spores are not produced until after the tree is dead.

6. It Can Confuse the Heck Out of Those Who Come Across It

I. glomeratus is a mushroom of many guises. It can be totally resupinate, sheeting a fallen log, or seriously pileate. Parts, or all of it, can be white, brown, orange-yellow, black, sulphur yellow, or grey. It can be covered with black guttation, or remnants of these drops, or it can have none. Its upper surface can be coated in golden or sulphur-coloured spores, or there can be no sign of yellow anywhere. Particularly when there are no spores or black guttation evident, it can leave people scratching their heads about its identity. The following selection of photos are all I. glomeratus.

Inonotus glomeratus yellow cap resupinate polypore

Inonotus glomeratus brown shelves polypore

Inonotus glomeratus

young Inonotus glomeratus

Log spray-painted yellow spores Inonotus glomeratus

Inonotus glomeratus yellow and white polypore
dried black exudiate
polypore yellow spores
white edge brown polypore Inonotus glomeratus

Monday, 11 July 2016

Unveiling the Veiled Polypore: Cryptoporus volvatus

dissected remains veiled polypore
I spent an hour this morning carefully destroying fifteen veiled polypores—totally ripped them apart. Odd thing to do? Nope, not in my neck of the woods.

Cryptoporus volvatus produces acorn- to walnut-sized cream-coloured bubbles on the bark of recently dead and usually still standing conifers. Sometimes it also appears on living trees. 

C. volvatus grows on recently dead or dying conifers.
At first glance, these cute little polypores might be mistaken for small, wood-inhabiting puffballs, or even the immature fruiting bodies of the slime molds Reticularia lycoperdon or Lycogala reticularia.

Lycoperdon pyriforme Lycogala flavofuscum Reticularia lycoperdon
Cryptoporus volvatus can superficially resemble (left to right)
Lycoperdon pyriforme puffballs (Wikipedia), or immature slime
molds such as Lycogala flavofuscum and Reticularia lycoperdon.

The reason they might not be immediately identifiable as polypores is that their pore surface is hidden, hence the "crypto" part of their name. Unlike other polypores, the spore-producing bottom side of the "veiled polypore" is covered by a volva-like flap of tissue—a covering that would apparently prevent the free movement of spores.  

Cryptoporus volvatus openings and lacquer-like coating
Note both the newly formed openings, or ostioles, and the
shedding of the lacquer-like coating (Dan Molter)
So how do these diminutive polypores manage to get their spores out into the larger world? For many years it was assumed that insects and other arthropods that are known to spend time inside these fungi are the obvious vector for spore dispersal. C. volvatus even provides a convenient doorway to gain access: a small ostiole that forms at the juncture with the tree, which enlarges as the spores mature. 
Veiled polypore ostiole or hole
The opening at the top of the photo is the ostiole that forms
in the pore covering. The other holes were made by insects.
But a 1980 study cast doubt on the insect link. The paper, published in Mycologia, posited that spore dispersal was primarily via air movement, just as it is for other polypores. The authors came to this conclusion by rigging up a bunch of C. volvatus with spore-collecting contraptions, then suspended the collected  spores evenly in a liquid, and finally counted them using a hemocytometer, a gauge invented in the 19th century for blood cells. And they found a lot of escaped spores—an average of 2.5 billion per basidiocarp. That's a lot of spores sent on their way by wind—comparable, in fact, to other polypores. 

So why evolve a covered pore surface in the first place? The authors suggest that it has to do with moisture retention. Many polypores are capable of resuming sporulation after they have completely dried out when they are rehydrated by rain. Not the short-lived C. volvatus, which usually grows well above the ground on standing trees where it is open to the desiccating effects of sun and wind. Basically, it's got only one chance to produce spores, so a protective tube covering makes sense. 

Cryptoporus volvatus brown shiny lacquer cracks
The lacquer-like coating that covers young C. volvatus
is also thought to help retain moisture.
But that air-movement and spore dispersal study didn't make the people who had a gut feeling that insects still belonged in the equation go away. Nope. This isn't surprising considering the number and variety of mini spelunkers that choose to hang out in dark, spore-covered, Crytoporus caves. 

Insects, like this one with metallic red elytra, tuck themselves
tightly into the corners when you remove the covering.
One Japanese study that looked at 438 separate C. volvatus basidiocarps counted a total of 8,990 individuals, insects that belonged to 17 different species, including 4 specialists that prefer C. volvatus over everything else. Some of these little mycophagists munch on the tubes, some eat the flesh, while others focus on the fallen spores. Predators enter as well. Several studies around the world have focussed on these insects and the possibility of their being a necessary vector for spore dispersal. 

This sporocarp has been completely hollowed out by insects.
In one Korean study, beetles were collected from C. volvatus basidiocarps and then "vortexed" (what a great verb!) in ethanol to dislodge any spores that were attached to them. Again, a hemocytometer was used to count the spores each insect specimen carried. And they carried a lot: from 10,000 to half a billion. That's a lot of spores to be moving around for no good reason. 

Basidiocarps of C. volvatus actually grow out of exit, entrance, and ventilation holes made by bark beetles—bark beetles that kill conifers but that are not known to feed on the fruitbodies of C. volvatus. So how do the spores gain entry to a tree? Air movement can't be enough. But it turns out that at least one species in the Korean study that feeds/breed inside the volval chambers of C. volvatus also spends time under the bark of dead conifers, so it could easily be a vector for spore transfer, perhaps to a passing bark beetle. 
My grand haul of Cryptoporus volvatus critters,
minus a bunch of tiny larvae.
I found this attractive, metallic beetle inside a
Cryptoporus volvatus "cave." Can anyone name it?
This guy was similar to, but a lot smaller than 
beetles I regularly find on Pleurotus species. 


T. C. Harrington, Release of Airborne Basidiospores from the Pouch Fungus, Cryptoporus volvatus, Mycologia, Vol. 72, No. 5 (Sep. - Oct., 1980), pp. 926-936

Kohmei Kadowaki, Species coexistence patterns in a mycophagous insect community inhabiting the wood-decaying bracket fungus Cryptoporus volvatus (Polyporaceae: Basidiomycota)Eur. J. Entomol. 107: 89–99, 2010
C. volvatus on E-Flora BC

Thursday, 5 May 2016

When Is Fuligo septica Purple? Not Very Often!

I was out with Ruby last summer when I spotted an unusual patch of colour in the distance, a pale, pretty, pinkish violet perched atop a moss-covered oak log. As Ruby and I got closer, it became obvious that it was not a wayward bit of plastic or a spent balloon, it was something special.
Daedaliopsis confragosa  and Hypoxylon fragiforme and dog
Ruby with Daedaleopsis confragosa
and Hypoxylon fragiforme
Now, Ruby is a fabulous foray companion — not only does she make me laugh, I can also count on her to give me warning of approaching wildlife, such as bears, when I’m silently concentrating on fungal marvels. Now that she’s outgrown her jubilant puppy habit of pouncing directly on top of whatever exciting specimen I happen to be focusing my camera on, she is almost perfect.

Ruby is a log-walker.
Almost, but not quite. Her one remaining flaw (besides, like most dogs, occasionally rolling in something nasty - see my last post), is that she is a log-walker. There’s nothing Ruby loves more than to scramble up onto a fallen tree and elegantly walk it from one end to the other, as sure-footed as a balance-beam gymnast — a problem if I’m trying to photograph or collect a specimen from said log.
I knew I had to distract her, so I gathered a few sticks and began throwing them until I reached my quarry. What I found was a large patch of not-quite-set, but fully-formedslime mold, Fuligo septica, but a F. septica that was, of all things, a beautiful lavender instead of bright or dull yellowish.

Immature violet Fuligo septica
Immature violet Fuligo septica 
I managed to keep Ruby at bay long enough to collect a sample and took it home. I set it aside to mature and produce spores and started reading. I took more pictures. I looked at under the microscope. I read some more.

pale pink plasmodium Fuligo septica
Immature violet Fuligo septica showing pale pink plasmodium 
I found a reference to a F. violacea(Persoon, C.H. 1801. Synopsis methodica fungorum), but little information about it other than the original Latin description which refers to a yellow outer shell. Mycobank clumps F. violacea together with numerous other synonyms of F. septica.  

Purple pink Fuligo septica mature
Violet Fuligo septica two days later
Having reached the limit of what I could find on line, I sent photos to Michael Warnock, one of the myxomycete aficionados of my local club, the Mycological Society of Toronto.  Michael replied with the suggestion that my specimen might be a F. septica that had been attacked by the myxomycophagist (slime mold eater), Nectriopsis violacea. I’ve actually found this curious character before. It looks completely different than what I’d found.

Nectriopsis violacea growing on Fuligo septica.
Nectriopsis violacea, an ascomycete, feeds on Fuligo septica.

dark purple perithecia of Nectriopsis violacea
The perithecia of Nectriopsis violacea are royal purple when moist.
N. violacea is an ascomycete that specifically targets F. septica. It’s not terribly common — I’ve only seen it twice, both times at the Cain Foray near Algonquin Park in Ontario — but it’s unmistakable. Its colour ranges from dark purple to purplish white (not pinkish lavender), peppered with minute darker pimples, the perithecia through which its spores are released. Not what I had found.

perithecium of Nectriopsis violacea and Fuligo septica spores
A squash-mounted perithecium of Nectriopsis violacea 
along with globose spores of Fuligo septica
F. septica, known for centuries as Flowers of Tan because of its proclivity to colonize the piles of tannin-filled wood chips used in the leather tanning industry, has several commonly found forms. The dull beige version that can sometimes produce massive fruitings on hay or straw or compost piles has been fondly called Dog Vomit Slime (self-explanatory). In Scandinavian folklore, it's identified as the vomit of troll cats, a creature I want to see. 
In my neck of the woods, I more often encounter a yellower, more compact version that matures from sometimes startlingly yellow, huge masses of plasmodium, a stage that gave rise to another descriptive moniker, Scrambled Egg Slime. This plasmodium sometimes matures into large, dull-yellowish amorphic patches on logs and trunks of trees. Other times it separates into much smaller, individual, deep cadmium yellow pillows. All of the above forms contain dark purplish brown masses of globose, minutely spiny spores that all look the same microscopically. None of these forms are pinkish violet. Not even close. But I did read that the final colour can have a connection to the acidity of the substrate on which its aethalia (pillow-shaped fruiting bodies) have chosen to form.

Yellow Fuligo septica plasmodium
A metre-wide patch of Fuligo septica plasmodium,
also known as Flowers of Tan
Under the microscope, I found another anomaly — the spores of mine all contained oil droplets of various sizes. So did I actually have a different species? Or did my sample have some kind of weird infection?

Spores of violet Fuligo septica showing oil droplets
Spores of violet Fuligo septica showing oil droplets
I found an old paper describing techniques for propagating F. septica and decided — in my completely amateur way — to try to grow mine. I put some spores on a sprinkling of ground-up oatmeal on a damp coffee filter in a covered container. Within a couple of days, the spores produced a flash of pigmented growth. The first day it was a bright, deep fuchsia, which then settled down to more of a pale pink, like the interior of the original immature aethalia. My experiment then stagnated and did nothing until it began to smell. I threw it out.

germination of spores produced pink fuchsia pigment
My spore-growing experiment produced brief pigmented growth.
Still, there had been no sign of the usual bright yellow pigment that F. septica commonly produces, so what did I have? I decided to go to the top of the myxomycete food chain and emailed Frederick W. Spiegel at the University of Arkansas, a specialist in Mycetozoans, a grouping of slime molds that include the genus Fuligo. He, in turn, forwarded my message and photos to his colleague, Steven L. Stephenson, author of numerous publications, including the indispensable Myxomycetes: A Handbook of Slime Molds, noting that Steven had likely seen more F. septica fruitings than any other human being.

 Fuligo septica plasmodium
Normal egg-yolk yellow Fuligo septica plasmodium
Steven, at first unable to open my image files, suggested once again that I had a found a F. septica specimen that was being consumed by N. violacea. Once he was actually able to see my photos, though, he informed me that he had once come across F. septica approaching the colour of mine, possibly in Alaska. He also said that oil droplets and other inclusions are found in a lot of myxos. His final word was that, as F. septica is almost certainly a species complex, what I had found could only be called that until such time that someone decides to make the effort to sequence a series of collections. As per his suggestion, I have frozen my collection for future study. Any takers?

Many thanks to Michael Warnock, Frederick W. Spiegel, and Steven L. Stephenson!

Three stages of a pale yellowish beige Fuligo septica
Three stages of a pale yellowish beige Fuligo septica
Typical yellow sporangia of Fuligo septica

The colour of Fuligo septica can vary enormously
The colour of Fuligo septica can vary enormously.

References & Resources

Steven L. Stephenson, Myxomycetes : a handbook of slime molds, Timber Press, 1994.

Patricia M. Scholes: Some Observations on the Cultivation, Fruiting and Germination of Fuligo septica, J . Gen. Microbiol. (1961), 29, 137-148 

Tom Volk's entertaining Fuligo septica page

Some fun Fuligo septica history from Historical and Literary Botany: Containing the Qualities, Anecdotes, and supperstitions, relative to trees, plants, and flowers, which are mentioned in sacred and profane history - Volume 3 By Eliza P. Reid 1826 

Great series of photos from Wayne’s World of the different stages of pale version of F. septica plasmodium growing on wood chip mulch

The Myxomyceticolous Species of Nectria: Gary J. Samuels Source: Mycologia, Vol. 65, No. 2 (Mar. - Apr., 1973), pp. 401-420

An amazing BLUE Fuligo!!!