Summary: Forget Kit Carson, General Custer, Billy the Kid and Geronimo. The real actors (good and bad) in the Wild West were made of spores and mycelium! And they didn’t need fictitious sidekicks like Silver, Tonto, Rin Tin Tin or Old Yeller—they rode on the wind, or had very real “pardners” with six A Mycologist’s Fervently Biased Account of legs and numbering in the millions! How the American Western Frontier Was Molded Collectively, they sustained by Spores and Mycelium or destroyed vast landscapes, brought salvation or ruin to entire Frank M. Dugan communities, deluded prophets USDA-ARS Western Regional Plant Introduction Station, and financiers, or provided 59 Johnson Hall, Washington State University, sustenance and joy. Fungi made Pullman, Washington 99164-6402 USA the Wild West, and are still shaping [email protected] the West’s ecological, economic and cultural destiny. even stored grain. Sometimes (rarely, after the Civil War, areas in the central Key Words: blister rust, cereal thank goodness) fungi can cause USA were still “frontier” (certainly the rusts, dung fungi, Entomophaga, disease in humans. Mycelium (plural, Dakotas, Nebraska, Kansas, Oklahoma, mushroom, mushroom poisoning, mycelia) is a bunch of thread-like stuff even parts of Texas and Minnesota). mushroom wars, pébrine, Puccinia, that comprises the “body” of a fungus My boyhood was in “wild” Montana, rots, Saccharomyces, smuts, (whether mushroom or mold) and with Dairy Queen and Safeway within sourdough, Sporormiella, Tilletia, fungi make a variety of spores (about a comfortable walking distance, but with Valley Fever, yeast hundred different kinds) which grow up wild-roaming elk nearby and buffalo to become more fungi. within an hour’s drive. Perhaps “frontier” is best defined as close proximity to What is (or was) “The Frontier?” aboriginal inhabitants resisting, by force What is (or was) “The West?” of arms, encroachments of farmers, Although fungi are difficult to define miners and ranchers. (Today, most What are spores and mycelium? precisely for the lay person, defining parties use lawyers, seldom bullets or Fungi make spores, each of which “The Frontier” and “The West” is just as arrows, to resolve issues of land, water, makes mycelium, which makes more challenging, probably because they’re or cultural sovereignty.) Alternatively, spores—right? Okay, but what are fungi? always changing! For Americans of “frontier” might mean far (in distance A formal and lengthy definition would colonial times, the Frontier could be or time) not just from amenities, but precisely define the word “Mycota” anything west of the eastern foothills from passable roads, medical care and and might quickly alienate most of my of the Appalachians. Even after law enforcement, in which case the readership. So, for those not already independence, “western frontier” might frontier persisted well into the twentieth well familiar with fungi, suffice it to say mean today’s Midwest (e.g., Indiana, century in many places. I’ve only been that examples of fungi (“Mycota” and Ohio, even western Pennsylvania). on horseback twice in my life, and I’ve stuff that looks similar) are mushrooms “American West” today generally never seen an arrow fired in anger, but (edible or otherwise) molds, and yeast; implies geography considerably west I’ve worn out several pairs of cowboy that fungi can cause rusts, smuts, leaf of the Mississippi (probably because boots, served on a fire lookout tower spots or cankers on plants, and that “westerns” vastly outnumber movies miles from the nearest road, listened many fungi can rot fruit, vegetables or about Hiawatha), but during and just to the sweet howl of the coyotes, and 6 FUNGI Volume 5:1 Spring 2012 consider myself from “the West”—so, symbolic—imagine how much poop I hope to be forgiven if I leave such came out of just one of those!), and terms as “the Frontier” and “the West” was conspicuously resurgent with the intentionally vague. introduction of cattle, whose occasional So why were fungi (however you concentrations created veritable define them), important to the West, poopscapes (Davis and Schafer, 2006). or the Frontier (however you define it)? The presence of abundant Sporormiella There are several, overlapping areas of spores has also been correlated with impact. Fungi directly altered landscapes overgrazing by livestock in the 1890s by attacks on vegetation, or maintained (Davis and Kolva, 1977). In addition to landscapes via nutrient recycling; they Sporormiella species, those in Podospora deteriorated produce and commercial are especially important on cattle dung products, or were used to make them; (Angel and Wicklow, 1975, with many they caused disease in crops or (more other species discussed). The poop- rarely) humans; and they were a source degrading capabilities of Sporormiella of sustenance, and have become “crops” species were early recognized (Fig. 3). in their own right. In fact, harvesting rights for morels are one of the few remaining issues which might be resolved “informally” with fists, or even Figure 1. When buffalo graze, revolvers, more frequently than with buffalo chips can accumulate lawyers. If you’re in the wrong place with quickly! (Photograph courtesy of a little bad timing, parts of the West Ray Wegner, www.raydw.com). can still get uncomfortably wild, and all because of some pricey mushrooms. This fungus (and several relatives) The following are some of the more live off the remnants of plant material important examples of the myriad ways in herbivore dung. High numbers that fungi molded (pun intended) the of spores of this dung-decomposing western frontier. fungus in sediments are used to infer the past presence of large numbers of Impacts on western grasslands grazing animals, be they cattle, buffalo Paddies of the past: Who cleaned up after the buffalo? In spite of all the “western” movies with scenes of deserts, sagebrush or chaparral, to many of us, “The West” Figure 3. An unsung hero: conjures images of vast grasslands and Sporormiella intermedia, poop herds of buffalo. The most conspicuous components of these grasslands were destroyer (from an early mycological grass (duh!), buffalo, and buffalo poop. treatise, Griffiths 1901). We know, more or less, where all the Of course, fungi are not the only grass went (it succumbed to the plow, organisms documented as decomposing and was replaced by grain—what was dung. Arthropods and bacteria are very left went under livestock or asphalt), important, too. But of the arthropods, and where the buffalo went (well, they those belonging to a minimum of mostly got shot and skinned, or just shot, eight families are actually feeding on and were replaced by cattle, sheep and dung fungi that previously colonized horses). But buffalo were around for a the paddies (Floate, 2011, on cattle long time. And for every mouthful of dung in Canada). Both arthropods and grass going into the front end of the Figure 2. Even with the buffalo in microorganisms are essential to efficient buffalo, there eventually was poop decline, buffalo chips remained poop removal (if either is lacking, poop coming out the back end, and the same so abundant they could be used persists—as was the case with cow was true for domesticated grazers (Figs. for fuel (Photograph courtesy of paddies in Australia before the right 1 and 2). So, long before they got shot, Chronicle of the Old West). beetles were introduced). Ecological why weren’t buffalo wading at least relationships between dung beetles, ankle deep in buffalo paddies? What or mammoths (Davis and Shafer, 2006; fungi and bacteria are complex, but happened to all the poop? It was mostly Kinney, 1996). Actually, Sporormiella fragmentation of the paddies by insects “eaten” by fungi! was probably most abundant in the age apparently boosts rates of decomposition Who were some of the primary of North American grazing megafauna (Lussenhop et al., 1980; Wu et al., 2011). doo-doo fighters of yore? Sporormiella! (of which the mammoth is the most The species of fungi colonizing dung FUNGI Volume 5:1 Spring 2012 7 have a picnic on a nice grassy meadow, beginning in earnest before World War I, remember that it’s thanks to the fungi persisted for some fifty years. Although that you aren’t picnicking on top of poop. enormous in scale, the effort was not particularly successful, and such control Fungal disasters of forest and farm as exists today is largely on the basis of White Pine Blister Rust producing pine trees that are genetically European settlers were not the rust-resistant. But for decades Ribes only invaders to alter entire Western plants were hand-pulled or poisoned ecosystems. White pine blister rust is, as with chemicals by workers, including the name implies, a disease of white pine high school boys, men in the Civilian and other closely related pines (Fig. 5). It Conservation Corps, war internees, was introduced into North America from college students and others. Figure 4. The old in and out: dung Europe in (or about) 1906, and had a The eradication program began in fungi life cycle. devastating effect on the growth of white the western states in 1919. In the inland pines, drastically reducing the utility of Pacific Northwest (the “Inland Empire”) are highly diverse, and apparently differ these trees which were highly prized for the number of men pulling Ribes plants by the type of dung which they prefer, timber production. In nature, the rust in 1934 was 11,000 (Maloy, 1997); over e.g. pronghorn antelope versus bison, cycles back and forth between the pine 8,700 laborers were employed in the etc. (Herrera et al., 2011). A detailed and an alternate host, gooseberry and work in 1936 (Benedict, 1981). “Large synopsis of the importance, ecology, and currant bushes (Ribes spp.). Ultimately, numbers of teachers and college students taxonomy of coprophilous fungi (many control efforts involved attempts at worked on the project, since the Ribes of which must pass through the digestive destroying the alternate host plant on season coincided with their summer systems of herbivores in order for spores “more than 20 million acres of private, vacation” (Benedict, 1981, see Figs.