Tom: That's about all there is! Oh, you forgot morels. That's what I did my thesis on.

Leon: Is that what you were initially interested in, in mushrooms?

Leon: I see. So this is already in graduate school.

Tom: Mmm hmm.

Leon: But you said that when you were an undergraduate, you already did work at Argonne.

Tom: I did. I did an undergraduate research project at Argonne National Lab near Chicago on revegetating coal spoils using some nitrogen-fixing legumes.

Leon: At strip mines?

Tom: Actually, this was the remnants of a coal-washing operation. They're left with these piles of stuff they call gob, which is very fine particulate coal, and the pH is around one or two [highly acidic]. Almost nothing will grow in it.

Leon: So you did have some interest in mycology as an undergraduate.

Tom: I was an undergraduate at the University of Ohio, in Athens. That's where I took my first undergraduate course, with Jim Cavender. Then I went to Madison for graduate school, where I started with the Schizophyllum project. I was actually doing fungal genetics, mostly. Not even real mycology! Whatever that means.

Leon: So where did you grow up?

Tom: Ohio. Northeast Ohio. Girard, Ohio.

Leon: Just a local boy who went to Ohio University...

Tom: Well, that was where I was raised.

Leon: What made you decide to go into mycology?

Tom: Ummm, it was interesting. I found out that I could find free food. Everybody likes free food. I was looking at some other ecology sort of things. I had offers from other schools to do plant pathology sorts of things.

Leon: But you just wanted to do straight mycology.

Tom: Yeah. Though you know, I was never doing taxonomy with morels. I was looking at the life cycle: the ecology (or at least the biology) of the organism.

Leon: That seems to be the trend in mycology since, well... in the early part of the century, anything in mycology was taxonomy: arguing about where things belonged, and naming new species. Now there's this big emphasis on the biology, and...

Tom: I don't know if it's really that different with all the molecular techniques: there's still a lot of taxonomy - or systematics, really: seeing how things are related to one another.

Leon: But back then, except for a few people, there wasn't anything else. I also call it new because it doesn't seem to be something that the field as a whole has really gotten a handle on how to do yet. When I see ecological studies, they're always evaluating their sampling technique at the same time that they're trying to get their results.

Tom: That's probably true. But that's still a pretty new sort of thing. And there are many problems with sampling for fungi, and the main one is that they're not always there. If you're sampling for redwoods, you can be pretty sure that the redwood that's there one year is going to be there the next one. But if you sample one week a year for a fungus, it may not be there the next year.

Leon: Well, there are these new techniques where you dig up the soil and test bits of it for DNA...

Tom: That's better, but even that is iffy. It may not be there in that same spot. It may be nearby, or... not there at all.

Leon: So they're just difficult to work with.

Tom: There are technical problems.

White morels, Morchella esculenta. Photo by John Denk.

Leon: I see. Now when you graduated from UW Madison, you didn't get a university job right away, you went to Forest Products, and you were working with Hal Burdsall there.

Tom: Right. My first job was to look at all of the fungi that were listed in the culture collection as Polyporus sp. [that's anything unidentified with pores and a cap], Poria sp. [anything with pores and no cap], Corticium sp. [anything without pores or teeth that just lays flat on wood], and figure out what they were. I had to go back to the herbarium specimens and key them all out.

Leon: Oh, I see! They weren't asking you to grow the cultures out to a fruiting body

Tom: No, no! I wasn't trying to grow them; but since we already had cultures, we needed to know what they were. And so I did that for... a while. And then I started to work on other things, eventually working on a project on the fungi of Alaska, where Hal... collected a lot of fungi and brought them home for me to identify.

Leon: That was nice!

Tom: I never got to go there, so although I'm first author on a 46 page paper on the fungi of Alaska, I didn't go there until this year, with NAMA's "exotic" foray.

Leon: I'm starting to see why you have so much more practice in identifying mushrooms than most other mycologists. So did having to identify all those polypores and crust fungi drive you out of Forest Products and into academia?

Tom: No, I wanted to teach, so I went into La Crosse [the University of Wisconsin at La Crosse, where Tom is now] when the job came up. I had always been teaching: even when I was in Forest Products. Well, let me go back before that. In graduate school, I was a teaching assistant for the first few years I was there, and then one day I came back from a weekend and they told me that the mycology professor, Bill Whittingham, had died and would I please teach mycology for the remaining 13 weeks of the semester.

Leon: And had you made the switch over to mycology at this point?

Tom: Well, I was always doing mycology, but it was the fungal genetics. And... so I taught the course, along with my major professor, Tom Leonard, and I taught mycology a couple more times before I finished graduate school. So I'd had a lot of teaching experience; I knew I liked it. Also, I taught for two semesters while I was at the Forest Products lab. I had a full-time job there and a half-time job at Madison in the Botany department. I had taught mycology once there, in 1984, and three times after that. So when the position became available at La Crosse I applied for it and.... eventually got it.

Tom: That was part of my job at the Forest Products lab. When I was done identifying the Poria sp.s, and the Polyporus sp.s, then there was money available for the monograph of Armillaria, and so I did that. But we haven't published that yet.

Leon: Oh!

Tom: [laughing] We published part of it, but not all of it.

Armillaria tabescens, the Ringless Honey Mushroom. Photo by John Denk.

Tom: Well, yes, the key. And we published a nomenclatural study where we looked at the two hundred sixty two species that were at one time in Armillaria and Armillariella and found that they belonged in 43 other genera.

Leon: We won't recap the highlights of that here.

Tom: No, we will not. We didn't do most of the transfers, but we kept track of who did what and when. And in the meantime, we described a new species of Armillaria that we had known existed but didn't know what it looked like.

Leon: Oh. So you had a culture, like all those Porias and other things, but you didn't have a preserved mushroom.

Tom: Right. We knew what the mycelium did in culture. It wouldn't mate with anything else, so we knew it was a separate species, but we didn't know what the fruiting body looked like.

Leon: And this is what became nabsnona.

Tom: Right. We named it Armillaria nabsnona, North American Biological Species Nine.

Leon: Are there any remaining to be named still?

Tom: There is: North American Biological Species Ten has not yet been named. I've never seen it. To my knowledge, anyway.

Leon: And that's what's holding up the project? Someone needs to find a fruiting body?

Tom: Mmm hmm. And it's not a very common species. It seems to be restricted to Idaho, and perhaps Washington and California. I actually went looking for ten right before the 1993 NAMA foray, at Fort Worden, Washington, and found nine instead.

Tom: I was only peripherally involved in Laetiporus, as it was getting started. I was only involved at the very beginning of it - in the middle I went to La Crosse, so I haven't worked much on it recently. I just keep track of it. Most of it has been done by Mark Banik at the Forest Products Lab, along with Hal Burdsall.

Chicken mushroom (Laetiporus sulphureus), photo by John Denk

Leon: Now, the other work I know like this is Greg's work on Laccaria. Laccaria, Laetiporus, Armillaria - they're all rather small genera, and clearly circumscribed.

Tom: Well, now they are!

Leon: Right! And that's my question: is this going to be a trend? Instead of working on these big genera like Clitocybe, are people going to separate smaller, clearer genera out and that's how things are going to get accomplished? Or is there always going to be a Clitocybe, and a Russula, with hundreds and hundreds of species...

Tom: There are people working on Russula - I know that Steve Miller at the University of Wyoming is working on Russula, trying to figure it out... along with lots of other people. There is some work being done on Amanita, but not all the sections of Amanita, for example. You've got to chip away at some of the big genera: work on sections of it at a time, or even subsections...

Leon: So what's the next project for you?

Tom: Oh, I have my finger in a lot of things. I teach mycology and medical mycology; those are the two big things. I had 22 students in Mycology this Fall, and I had thirty-eight students in medical mycology last Spring, which they tell me is the largest medical mycology course in North America. I don't know if that's true, but it sounds good! I spend a lot of time teaching those courses, and every other year we do Plant-Microbe interactions. Seven of us do; well, four of us do it now..

Leon: Four professors.

Tom: Yup.

Leon: And that's even though fungi aren't microbes.

Tom: Oh, they're microbes.

Leon: You mean yeasts?

Tom: No, they're microbes, they're microscopic. That's all that matters. They don't have to be single-celled, that's different. Microbes can be... anything that's... mostly microscopic. Most fungi are mostly microscopic.

Leon: Okay, so you're talking about like the Glomales, and things like that.

Tom: Not just them; we do the all the fungi, even the ones that form big fruiting bodies, and all the plant pathogens, and.. mycorrhizae... fungi are microbes. We include them in our course, anyway, all with the bacteria and viruses and nematodes and, mycoplasmas, and things like that.

Leon: Nematodes are microbes also?

Tom: Well, they are for our purposes; they're small things. So then, every spring now I'm teaching Latin/Greek for scientists. Next time is my third time teaching it. And then I have lots of different research projects, mostly driven by what my students would like to do. I have more students than ever before, so now I'm busier than ever before. My first student, Marsha Harbin, worked on morel mycorrhizae. She won a graduate student award for our poster presentation at the 1999 meeting of the International Botanical Congress and the Mycological Society of America, in St. Louis. The next student, Sean Westmoreland, has been working on Hydnellum, looking at pigments for use in taxonomy.

Leon: And these are masters students?

Tom: Masters. We don't have a doctorate program.

And Nik Zitomer is working on antibiotics from fungal fruiting bodies, and Josh Burgess is working a molecular probe to look for Blastomyces in soil, to find out what its ecological niche really is.

I've a new student, Bridget Pfaff, who just started. She's going to look at indoor air quality, that sort of thing. Adam Gusse is working on the fungi of Alaska, and just spent the summer there. He used to be an undergrad here, but now he's one of my grad students. And.. I have several undergraduate students who are working on lots of interesting things.

Bernadette O'Reilly is working on some of the follow-up on the Entoloma-Armillaria story that I did with Dan Czederpilz;

Laura Appleman is looking at biocontrol of plant pathogenic nematodes using fungi; Nicole Aubart & Chau Nguyen are working with Nik Zitomer; Matt Schwenke is working with Adam Gusse on that project; and Maria Lee is working on malaria.

Leon: So, mostly now, you're an educator: at the web and your job, through these research projects.

Tom: Yeah, I let my students kinda pick what they like, within reason. Something that I can help them with, anyway. Mostly, the public needs to be educated about the importance of fungi, and teachers need to be educated about the importance of teaching about fungi, in their high schools and elementary schools. I've got the website, and I get email from high school and grade school teachers all the time. And grade school and high school kids. But then I ask my undergraduates how many of their teachers used the F word [that's "fungi"; both Tom and my proofreader insisted that I point that out to you!] in high school, and usually it's not very many.

Leon: Do you see a change in why people are getting into mycology?

Tom: Well, the field is becoming more diverse; it's not just working with fruiting bodies anymore. It's becoming obvious that there are different sorts of things that you can do with mycology - indoor air quality is an example. There are not many people who are very good at identifying molds, so they're in demand right now, because it's suddenly a big issue in the building industry. So there're a big rush to do that. There are lots of people who're excited about the molecular techniques and getting into that because it's interesting. There are always people who are interested in natural things, and amateur mycologists who become interested because of the eating aspect, which is fine with me!

Leon: So you think that it's mostly the non-professionals who are interested in the eating aspect?

For a long time, everyone thought that honey mushrooms parasitized Entoloma abortivum, but Tom and Dan showed that it's the other way around.

Tom: Not really. I mean, Peck ate most of his type specimens.

Leon: What? Really? So we don't have them any more?

Tom: Yeah, we don't have a lot of his types, because he ate them!

Leon: As a test? To see if they were poisonous or not?

Tom: Oh, I don't know why.

Leon: Wow. That would cause all sorts of problems...

Tom: Yes, it has caused all sorts of problems. Now we don't know what a lot of his species are.

Leon: What do you think are the big problems in mycology today? Where are the big questions to be figured out?

Tom: Well, there's a lot of things. A whole lot of ecological things, a whole lot of systematic things are big questions. Mostly, though, there's a lot of practical research to be done. There's lots we don't know how about how fungi interact with most other organisms. We don't know - - we're learning, but we don't really know - - about the phylogeny of the fungi: what they evolved from and how they changed and which groups came first. We know in general what probably happened, but ... there's a lot of details to be worked out that are difficult because there's no fossil record.

Leon: Well, there are some fossils...

Tom: Yeah, about four! [laughter] There's barely any fossil record except for spores.

Leon: Anyone ever tried to germinate those?

Tom: Well, they're fossil spores; they're stone, so...

Leon: Oh, duh.

Tom: You could try! They have germinated spores they found in glacial ice, you know, ten thousand year old glacial ice, so those do germinate.

Leon: And what were they?

Tom: I don't remember. whether they're molds, or... I don't remember

Leon: A future article for Mushroom the Journal!

Leon: I was told you predicted for the future a little machine where you could chop off [Tom is already laughing] a little bit of the fungus and it would analyze the DNA instantly and it would tell you instantly what it was.

Tom: Yeah.

Leon: Is that serious?

Tom: Yeah. I think it'll happen some day, yeah. But the problem is we don't have enough information to put in there, to know how to program it. Although you can make some significant advances using molecular biology techniques, someone still has to look at that fruiting body and figure out what it is in the first place, so you can match that information with its DNA. And we still don't know what 95% of the species are; they're not described to science.

When you name a new species, you're supposed to keep the original specimen that you based the species on. This original specimen is called the "type" specimen.

That way, when people think they've found another but they're not sure, they can check their find against the original. It also helps if you've forgotten to mention some crucial feature: later scientists can go back to the type specimen to check for that feature. If the type specimen isn't there, well...

Leon: That's macroscopic, or all of the fungi?

Tom: All of the fungi.

Leon: I assume that with macroscopic fungi the percentage is a little better.

Tom: It's probably better. I haven't seen data on it.

Leon: Yeah, I mean, we find about half a dozen per year in Chicago that we can't identify, and probably three or four of those are undescribed. The rest were things that are known from some paper somewhere, but aren't in the field guides... Oh, talk about your field guide! What are you going to put in your field guide?

Tom: Ooohh, I'll have one some day.

Leon: Is it going to include all these species that are known but aren't in any of the other field guides?

Tom: Mmmm, it'll have a lot of species, anyway. It'll have pictures of around 1100 species in color. and then there'll be a lot of pictures on the CD that goes with it. There'll be about 4000 pictures on the CD, I figure.

Leon: And these are multiple pictures of the same thing?

Tom: There may be multiple pictures of the same thing where there's a variant. You know, where sometimes it's white, sometimes it's yellow, the other color-form will be on the CD. And you know sometimes you see in a field guide, where it says something like "also similar is such-and-such, except that it has... blue teeth." There'll be a picture of that in there. On the CD. The book itself will have the pictures on the same pages as the descriptions, dichotomous keys and lots of introductory material based on the families.

Leon: So you've decided which families you're going to use.

Tom: Yeah, I've got a pretty good idea.

Leon: Regardless of whether they change in the meantime?

Tom: Well, the problem is how do you reconcile the molecular biology with that. In a field guide, you're still going to need the things with teeth keying out together, because that's how you have to do it, macroscopically, and so we'll make comments on where these belong. Evolutionarily, genetically.

Leon: So it's going to have artificial keys?

Tom: Yeah, a key has to be artificial.

Leon: Well, it's going to be interesting to see how it turns out.

Tom: Yeah, it sure will be! No, really, I do have some idea of what it's going to look like.

Leon: Are you collecting pictures?

Tom: Yeah. I have a lot of 'em, and I have offers from a lot of people; that's one of my projects.

Leon: So it's going to have a lot of previously un-field-guided species.

Tom: Probably. There will be more polypores and crust fungi than usual.

Leon: It seems like there's a two-fold educational challenge that's facing you. One is educating the general public about fungi in general, and the other one is educating the greater mycological community about things like... crust fungi

Tom: Right.

Leon: Things that aren't perhaps perceived as exciting.

Tom: Things that are perhaps obscure, but are very important: crust fungi are doing all sorts of decomposition in the forest. I mean, there's ... oh, ten people in the world who can identify them with any kind of consistency or competency. No, probably not ten. In fact, not many more than five.

Leon: So, you did the thesis on the morels, and then you went right into the job with Forest Products?

Tom: Well, I spent a year and a few months in the same lab I'd been in, with Tom Leonard at Madison.

Leon: Still working on the morels?

Tom: I was writing up my papers and doing some further work. Cultivation - cultivation of other fungi - and that sort of thing. We were actually working with a shiitake company in Madison at that time, and that's a whole big long story.

Leon: Do you do you have any opinions on where fungal cultivation is going to go? Do you see Marsha's work [on morel mycorrhizae, discussed above] leading to any advance in morel cultivation?

Tom: Could be. I don't know. I don't... There's going to be a breakthrough somewhere, but I don't know where it's going to be. Marsha's work just shows why it's difficult to cultivate morels; it doesn't immediately suggest a way to do it.

Leon: Like planting a little tree in the tray with the morels.

Tom: Nooo. My father always said "Why don't you just plant dead elms?"

Leon: It sounds, from the work with the blewits you mentioned, that plain old nutrition is still something of a mystery.

Tom: Yeah, we just played with that a little bit, but there was definitely something it needed that it wasn't getting. We had a project to work on mushrooms from beer waste, just to evaluate the substrate for possible economic use. We tried the easy stuff, and it worked okay; about as well as the initial attempts with sawdust and shiitake logs, so it just needs some development.

Leon: For the easy stuff.

Tom: Yeah.

Leon: And the blewits are just...

Tom: We just don't know what's going on. They're secondary decomposers, just like Agaricus, so there's a whole ecosystem there that it's getting something from, and we don't know what that is. Agaricus is actually really hard to grow also - it's very complicated - but we know the method now. When you know the method, then it's easy. But Agaricus is difficult just to culture: you put it in agar on a petri dish and it doesn't like to grow there.

Leon: Because it needs what?

Tom: Something that we don't know. Something that it gets from the compost that it doesn't get from agar.

Leon: It could be other living things that it attacks, it could be nutrients...

Tom: It's probably nutritional. Normally, the Agaricus is grown in a mishmash of other organisms, and we don't even know what most of them are.

Leon: I remember reading that it wouldn't fruit in sterile compost; that it needed these bacteria...

Tom: Right, something is stimulating them, but we don't know what it is.

Leon: So do you think we're going to figure out how to cultivate mycorrhizal fungi?

Tom: I don't know. That's a whole new level of difficulty. I don't think that most mycorrhizal fungi will ever be cultivated without some major breakthrough. And if I knew what that was, I...

Leon: You'd be working on it.

Tom: I'd be doing it. I'd be... sitting in the lap of luxury.

Leon: Yeah. Well, it's funny to think about morels being mycorrhizal. I don't really know of any other mycorrhizal ascomycetes.

Tom: Truffles.

Leon: Oh. Duh.

Tom: Rhizina undulata. There's a lot of mycorrhizal ascomycetes. Elaphomyces. Quite a lot of them. Most of them are below ground, of course. But there're probably a lot of fungi that are mycorrhizal that we don't recognize as mycorrhizal, because of different strategies of fruiting. Like fruiting after the tree is dead, the way morels do. There are probably a lot of other things that are doing that, but we don't recognize them.

Leon: Like the Coniophora we found at the walk this morning, which grows up on the bark of the tree and then fruits.

Tom: Right, it's a bolete. It's related to Suillus.

Leon: That's so weird. So it grows mycorrhizally in the soil, and then sends this big slimy thing up the bark to fruit. [Check Coniophora out in your field guides and tell me whether you would have guessed in a million years that this thing is related to boletes.]

Tom: Right. Most fungi have to form this large fruiting body to raise themselves up above the ground so that their spores can be dispersed more easily. Well, this doesn't; it just uses the tree for the same purpose. So it's probably very advanced. It's probably not a primitive character.

Leon: So there may be other fungi like the Coniophora that do something that we don't recognize,

Tom: Yeah. There's some misconception that the name and the relationships that the name implies are the end product of everything. But the name is not the end product, the name is just the beginning of learning everything else about the organism, because once you can learn to identify it, then you can find out everything else that's interesting about it. The name of it is not interesting. The name is not interesting at all...

Leon: Like then you can start saying things like "[Armillaria] gallica is the honey mushroom that grows in troops on the ground" and...

Tom: ...and it's not a pathogen, or at least it's a very weak pathogen. That's how you learn things about the organism. They say that Taxonomy was the first science. "What is it? Can I eat it?" You have to figure out what things are in order to work on them - to do the really interesting stuff. That's how we use the molecular biology: it's telling us that these organisms are linked, and now we can go back and look at why. What are the characters that are important - that tell us that these things are related? Because for a lot of these things, it's how you look at the features to decide what name it belongs under. That's a lot of what we need to learn from these people: which characters are important.

Leon: Macroscopically.

Tom: Macro, micro, or any other way we can do it: biological species, mating, culturing, whatever else we can do to separate 'em. So that to me is the great value of the molecular techniques: so we can find out all the other interesting things about them.