Tag: mushrooms

Deer Truffle

Deer truffles look like small clods of dirt; sectioning reveals spores. Note acorns for size.

Common Name: Deer Truffle, Deer balls, Hart’s balls, Warty deer truffle, Fungus cervinus (cervus is Latin for deer), Lycoperdon nuts (Lycoperdon is a genus of puffball fungi) –  Truffle is a French variant of the Latin word tuber meaning lump or knob. Both truffles and tubers (like potatoes) are generally globular in shape. The association with deer is attributed to finding them in locations frequented by stags during mating season. This gave rise to the belief that truffles are aphrodisiacs.

Scientific Name: Elaphomyces granulata – The generic name is a literal translation of Greek,  deer (elaphos) fungus (mykes). The Latin granulum is used directly in English as granule, referring here to the protuberances on an otherwise smooth surface. A loose translation of the scientific name would be “warty deer fungus,” one of the common names.

Potpourri: The deer truffle genus Elaphomyces is one of the most important mycorrhizal genera in temperate and subarctic forests, establishing and maintaining the ecosystem balance between plants and fungi. They are also an important source of food for small mammals like mice and voles on every continent except Antarctica. Deer truffles are equally favored by their namesake, notably the red, roe, and fallow deer species of Europe. Of the 49 species of deer truffle so far recognized worldwide, 20 are European. E. granulatus is one of the  most important North American species. A related species, E. muricatus, has been used in Mexico, both as “a stimulant, for remaining young and treating serious wounds” and “in shamanic practices in association with psychoactive Psilocybe species.” [1] Limited research in the 21st century has revealed that E. granulatus has enzymes that are known to reduce inflammation in addition to a variety of anti-oxidants with potential medicinal applications for humans.

Deer truffes are among the most common of underground fungi globally, and equally one of the least documented. The lack of scientific research on deer truffles is due partly to their sub rosa, subterranean obscurity and ignorance about their ecological importance. Even when uncovered, they look like lumpy balls of dirt. However, unlike the more famous black and white truffles of Europe, they are neither redolent with beguiling aromas nor palatable. Taste testers report that the main body is like “thick cream that tastes like nothing,” a rind that is “rubbery but can be chewed quickly,” and “a taste that goes in the direction of earthy forest floor.” They are, nonetheless, relished by rodents. [2]

Truffle is defined as “any of an order (Tuberales) of fleshy, edible, potato-shaped ascomycetous fungi that grow underground.” However, truffle is broadly applied to any hypogenous (below ground) fungus that is shaped like a tuber, which is the thickened part of an underground plant stem like those of the yam, cassava, and potato. According to historical etymology, any roundish shaped globule dug out of the ground was a tuber and/or truffle. [3] The distinction between plant and fungi kingdoms was not established until the 20th century so it would have made no difference whether the earthen globule was a plant tuber or a fungal truffle. The lumpiness meaning is inherent in chocolate truffle, a confection shaped like a truffle having no fungal ingredients. The terms edible and ascomycetous in the definition require some elucidation.

Edible does not necessarily mean by humans, but merely that it is or can be eaten for nutrition by an animal. Being edible is also a matter of importance, as fungal truffles reproduce by spores that must be transported for propagation. Above ground or epigenous fungi/mushrooms accomplish this with airborne wind dispersion, a mechanism not available to truffles buried several centimeters deep. Truffles must usually be consumed by an animal to transport the spores to new fertile ground and must thus be at least palatable. The need to attract animals is key to the inimitable smell and taste of certain species of truffles. It is probable that some truffles are unearthed and broken open without being eaten to release spores, so consumption is not absolutely mandatory although certainly the norm. Insects and worms that tend to feed on fungi may also play a role. Edible is a broad term in this context.

The term ascomycetous is a bit more complicated. The vast majority of fungi typically called mushrooms are in the subkingdom Dikarya which means “two nuclei” in Greek. Dikaryotic cells replicate with cell division of one nucleus from one “parent” and one from the other as they grow so that each new cell has two nuclei until the creation of reproductive spores through meiosis to pass the combined DNA to future generations. The way in which spores are produced divides Dikarya into two phyla, Basidiomycota and Ascomycota. Basidiomycetes produce four spores at the end of a club-shaped structure called a basidium. Most of the fungi that look like a mushroom with a cap or pileus at the top of a stem or stipe in addition to the various bracket fungi, puffballs, and stinkhorns fall into this category. Ascomycetes produce eight spores inside a sac-like structure called an ascus, Greek for wineskin or bladder. The asci are typically arrayed on a concave surface giving rise to the more common name “cup fungi” for ascomycetes. Most fungi, including yeasts, rusts, smuts, lichens, and, notably, truffles are ascomycetes. [4]  False truffles are basidiomycetes that look like truffles―ball-shaped structures that grow below ground.

Both truffles and false truffles followed different ancestral trajectories to become nearly identical in size, shape, and disposition due to similar environmental factors, a process called convergent evolution. Richard Dawkins offers that this is because “however many ways there may be of being alive, it is certain that there are vastly more ways of being dead.” Organisms tend to come up with similar ways to survive in the unforgiving environments of nature. Life above ground can be dangerous due to predatory and environmental challenges making it  advantageous to seek refuge in the soil. Many animals also do this. It is hypothesized that truffles evolved from cup fungi and false truffles evolved from mushrooms like agarics and boletes as a matter of random mutation resulting in improved survival. However, it could equally be the other way around, i.e. fungi may have originally been underground ”truffles” and evolved mushroom stems and gills for spore wind dispersion. DNA sequencing of the world-renowned Périgord black truffle corroborated the estimate that Pezizomycetes, the largest group of Ascomycota that includes truffles, separated from other fungal lineages 450 million years ago, just as the first plants advanced onto land from the sea. [5]

Deer truffles from Germany. Note root-like attachment to the mycelium.

Most fungi start as a root-like structure called a hypha emanating from one spore joining up with another hypha from another spore to  form a mycelium, the tangled mass of hyphae that defines the fungus. Since no species can survive without reproducing at some point, the mycelium must somehow send spores somewhere to start anew. Just as plants have devised ingenious ways to spread seeds, so have fungi to spread spores. Mushrooms start as underground bodies called primordia that are formed by the mycelium. They erupt upward on a stem into open air when the time is right to expose the spore bearing gill or pore surface to transporting winds. In the case of truffles and false truffles, the spores are contained in the tuber-like body that is attached to and grows from the mycelium but remains underground. The evolutionary pathway for the truffles and false truffles was to attract animals with enticing smells, not all that different from plants producing flowers with complex chemical scents to attract pollinators. Note that it is important for truffle smell signaling to start only when the spores are fully mature and ready to transport. Animals drawn by the smell to eat them transport truffle spores unwittingly wherever and whenever they “go.” [6]

Animals attracted to truffles and false truffles are globally diverse, inclusive of  deer, bears, and rabbits in the Northern Hemisphere and armadillos, baboons, and wallabies in the Southern Hemisphere. [7]  Underground fungi offer a food source that is relatively independent of surface conditions making them especially important to cohabitating animals. While most if not all forest dwelling mammals consume truffles on occasion, it is the burrowing squirrels and voles that are best equipped to use them as a major food source. With a keen sense of smell and claws to dig up buried acorns, there can be no doubt that squirrels are truffle aficionados. One well studied example is the California red-backed vole of the Pacific Northwest which subsists almost entirely on truffles. A study in the Oregon Coast Range involving vole capture and evisceration found that truffles made up 85 percent of consumed food, the balance was mostly lichens, also predominantly fungal. The northern flying squirrel, with a range from Alaska to North Carolina, is a nationwide spreader of truffle spores. [8] The extent of the role that truffles play in forest ecology as both providers of key soil nutrients like phosphorous and nitrogen to trees and as food for foragers is not well studied and therefore mostly unknown. This relationship is called mycorrhizal (meaning fungus root in Greek) and was first discovered by a biologist named Albert Frank in 1885 while employed by the King of Prussia to attempt to cultivate truffles. [9] Since there is no above ground evidence and animals need to be literally caught in the act, data are mostly anecdotal. However, one can gather some insight of the range, diversity, and importance of truffles from the aptly named “desert truffles.”

A desert is a dry, barren place incapable of supporting almost any plant or animal life. And yet, truffles thrive across North Africa and the Middle East all the way to China. Eking out a tenuous existence with shrubby plants with which they are mycorrhizal, they are surprisingly ubiquitous. They are sold in many local markets and consumed as an important food source over a vast region, noted for having a taste characterized as “delicate, not pungent.” They are reportedly relatively easy to find as they grow close to the surface and make the ground harder, a property that can be discerned with experience by rubbing a bare toe over the area. [10]  As Mesopotamia was the cradle of western civilization, the long history of truffles as both food and medicine there is telling. Truffles have historically been a substitute for meat throughout the Arabian peninsula. Truffles (kama’ah in Arabic) appear in the Koran as preventive medicine, used as promoters of longevity and good health much as many other fungi are in Asia. This a measure of their reputed anti-oxidant, anti-inflammatory, and immune modulating activity. 11] The cultural importance and extensive range of desert truffles across a broad swath of Eurasia is a strong indicator that they are key components of the plant-fungi global ecological partnership. While truffles are surely common and keystone in many regions, almost  all of what is known and studied about the nature and nurture  of truffles derives almost in entirety from detailed study of a few species that are among those granted the rubric “true truffles.”

True truffles are the epitome of European gastronomy. The black truffle of the Périgord region in southern France (Tuber melanosporum) is surpassed only by the white truffle of the Piedmont region of northern Italy (Tuber magnatum) in desirability and exorbitant cost. The reason for the difference is supply and demand, the universal economic law. White truffles are rarer because, unlike their cultivated French cousins, they grow only under naturally appropriate conditions and require specialized skills to locate. Consequently, in local Italian trattoria, one can purchase risotto with black truffles for about 20 euros, but risotto with white truffles will run over five times as much. [12] The reason for truffle demand is the redolence they impart to food, beguiling gourmands in their search for epicurean nirvana. It is telling that truffles were originally hunted with domesticated female pigs attracted by their aroma which includes the steroid alpha-androstenol, also found in the saliva (and breath) of rutting boars. The same chemical is found in the underarm emanations of men and in the urine of women, and, while the sexual role of the steroid in human sexuality has not been proven, it has been demonstrated. Men rating photographs of (clothed) women for sexual attractiveness gave higher marks when smelling alpha-androstenol.  [13] In that smell is intertwined with taste according to the neural-networked brain, the irresistible allure of  truffles to humans probably has deeper meaning and possibly including subliminal sexual arousal. It is no wonder that they are considered to be aphrodisiacs. Perhaps at least mentally they are.

It is almost certain that boars that have roamed wild across Europe for millennia were the coevolutionary partners of white and black truffles, spreading their spores far and wide. It is probable that humans first became aware of truffles in association with hunting wild boars. Thus began the long partnership between domesticated pigs and people in the pursuit of pleasure. Dogs have mostly replaced pigs as the truffle hunter’s sensory companion. Heavy, sedentary pigs required carting to truffle forest habitats and had to be forcibly prevented from eating their quarry; many a truffle hunter lost a finger to an overzealous pig. Dogs are not sexually attracted to truffles and must therefore receive olfactory training, much like drug-sniffing dogs of the DEA. This takes a great deal of time and effort, which must of necessity include the use of valuable, short-lived truffles. Trained truffle dogs are dear, commanding prices of over 15,000 euros but rarely sold. They can transit and search woodlands with ease and are not overwhelmed by lust for consumption. In fact, most truffle dogs don’t even like them, though apparently some do. Dogs have different taste preferences, as do their best friends. But not pigs, apparently. [14] The wild boar fungus story has a recently discovered twist. Of 48 boars killed in hunts in Bavaria, Germany, 88 percent had radioactive cesium levels (from Chernobyl) exceeding safety standards. It is considered likely that eating fungi that tend to bioaccumulate heavy metals were the source, especially truffles. [15]

References:

1. Paz, A. et al . “The genus Elaphomyces (Ascomycota, Eurotiales): a ribosomal DNA-based phylogeny and revised systematics of European ‘deer truffles'”. Persoonia. 30 June 2017. Volume 38 Number 1 pp 197–239.

2. “Deer Truffles – biology, ecology, distribution and occurrence of Elaphomyces or False truffle” https://www.umweltanalysen.com/en/elaphomyces-deer-truffles/  

3. Neufeldt. V. ed Webster’s New World Dictionary of American English, Third College Edition, Simon and Schuster, New York, 1988, p 1435, 1438.

4. Lincoff, G. National Audubon Society Field Guide to North American Mushrooms, Alfred A. Knopf, New York, 1981, pp 323, 377.

5. Martin, F. et al “Périgord black truffle genome uncovers evolutionary origins and mechanisms of symbiosis” Nature, 28 March 2010, Volume 464 pp 1033-1038.  https://www.nature.com/articles/nature08867 

6. Arora, D. Mushrooms Demystified, Second Edition, Ten Speed Press, Berkeley, California, 1986 pp 739-741, 841-865.

7. Trappe J. and Claridge A.” The Hidden Life of Truffles” Scientific American April 2010.

8. Stephenson, S. The Kingdom Fungi, Timber Press, Portland, Oregon, 2010 pp 200-205.

9. Frank, A.B. “Über die auf Wurzelsymbiose beruhende Ernährung gewisser Bäume durch unterirdische Pilze” [On the nourishing, via root symbiosis, of certain trees by underground fungi]. Berichte der Deutschen Botanischen Gesellschaft. 1885 Volume 3: pp 128–145.

10. Schaechter. E. In the Company of Mushrooms, Harvard University Press, Cambridge, Massachusetts, 1997, pp 161-167.

11. Khalifa, S. et al “Truffles: From Islamic culture to chemistry, pharmacology, and food trends in recent times”  Trends in Science and Food Technology, Volume 91, September 2019, pp 193-218. https://www.sciencedirect.com/science/article/abs/pii/S0924224418303406 

12. Goldhor, S. “Hunting the White Truffle” Fungi. Volume 8 Number 3, Fall 2015, pp 18-23.

13. Kendrick, B. The Fifth Kingdom, Third Edition, Focus Publishing, Newburyport, Massachusetts, 2000 pp 281-283.

14. Campbell, D. “Sketches from the Italian Truffle Hunt.” Fungi, Volume 11 Number 1, Spring 2018, pp 20-25.

15. Rains, M. “Germany’s radioactive boars are a bristly reminder of nuclear fallout” Science, 30 August 2023.

Destroying Angel – Amanita bisporigera

The key features of the Destroying Angel are the cup-like volva at the base of the stem, the stark whiteness of the stem, cap, and gills, and the partial veil hanging from the top of the stem just below the gills under the cap.

Common Name: Destroying Angel, Fool’s Mushroom, Death Angel, White Death Cap – The virginal whiteness of all parts of the mushroom are aptly described as angelic – beautiful, good, and innocent. The fact that it is anything but is conveyed by the addition of destroying with death-dealing toxicity.

Scientific Name: Amanita bisporigera – The generic name is taken directly from the Greek word amanitai, probably from the Amanus Mountains of southern Turkey where the noted Greek physician Galen may first have been identified the archetype, Amanita. [1] The specific name indicates that there are only two spores on each of its basidia in contrast to the four spores of other basidiomycete fungi. Virtually indistinguishable from Amanita virosa and Amanita verna which both frequently appear as synonyms in mushroom field guides.

Potpourri:  The destroying angel is a toadstool nonpareil. While the origin of the term toadstool is obscure, it cannot be a coincidence that tode stuhl means death chair in German, the language of the Saxons who emigrated to England. Its notoriety is not only because it is one of several mushrooms that contain deadly poisons called amatoxins, but also due to its close resemblance to Agaricus campestris, the edible field mushroom which is the cousin of the cultivated white button mushroom of supermarkets and salad bars. Both are white, similar in size and shape, and grow in the same habitat, primarily grass under or near trees. The destroying angel is the most dangerous of the numerous doppelgänger mushrooms as the deadly twin of a well-known and often consumed edible.  Misidentification absent knowledge of the subtle physical differences between the two can result in discovering the profound physiological differences with sometimes deadly result. The field white mushroom is nourishing. The angelic white mushroom is Shiva.

The cup at the bottom of the stem is the volva, the bottom half of the universal veil.

The key features that distinguish the destroying angel from similar mushrooms are straightforward if you know what to look for. First and foremost is the volva, (Latin for a covering like a husk or shell) which is the cuplike structure at the base of and surrounding the stem or stipe. The volva is frequently hypogeal, i. e. underground and out of sight. This means that it can only be positively identified by digging up the soil around the base of the mushroom. [2] However, it is the standard and preferred practice among mushroom gatherers to use a knife to cut through the stem cleanly at the base. This is done so the mycelium of the fungus from which the fruiting body mushroom grows is not seriously disturbed. The procedure is analogous to gathering apples from an apple tree. The fungal mycelium and the apple tree survive to produce new mushroom spores and fruit seeds for future generations. Using the standard harvesting technique, it is easy to see how the below the cut volva would not be noted.  White mushrooms must be dug out to the roots to avoid the dilemma of the death mushroom.

The only way to be certain that you have a puffball and not a Destroying Angel is to cut it in half.

The volva is the bottom part of what is known as a universal veil, a thin membrane that envelops the mushroom during the subterranean growth phase to protect the gills and the spores they hold from damage. The universal veil is a characteristic of all mushrooms in the Amanita Family. While there are a few other mushrooms that have a universal veil and its volva (such as the genus Volvariella named for this characteristic feature), it is a reliable identification feature for the destroying angel. All spore-bearing mushrooms are produced by the fungal mycelium underground as an ovoid called a primordium. Once they mature and environmental conditions are promising (like after rain) the extension of the stem causes the universal veil to tear around its circumference to expose the cap and gills of the fruiting body for spore dispersal. The volva is the lower part of the “eggshell” that remains attached to the bottom of the stem. Prior to upward extension, the destroying angel looks like a white egg, similar in appearance to a puffball, another type of edible fungus with which the destroying angel can be confused.  Some field guides include a picture of it in the puffball section to emphasize the danger of mistaken identity. [3] The only way to be absolutely sure is to cut the fungus lengthwise to reveal a cap and gills within.

Many mushrooms have what is known as a partial veil which also helps prevent damage to the reproductive gill surface. It is partial in that it only covers the underside of the cap, extending from the edges of the cap to the stem. When the mushroom cap expands fully, the partial veil also tears, in many cases leaving some remnants around the edges and a ring called an annulus attached to the stem just below the cap. In some cases, the partial veil remnant can be seen hanging like a draped clerical mozetta at the top of the stem. However, this annular ring is not well connected, and in many mushrooms with partial veils, there is no remnant. Most Amanita family mushrooms have both universal veils and partial veils with both a volva at the bottom and a ring around the stem as is the case with the destroying angel. The double protection afforded to the gills must have evolved due to the success of the species in propagation. Amanitas are one of the most prolific of all mushroom families. Partial veils and the remnant annulus are also a characteristic of the Agaricus family, which includes the edible field mushroom Agaricus campestris. They do not have universal veils with the tell tale volva.

The second prominent feature of the destroying angel is the stark whiteness of the cap, stem, and gills that has been described as having a “strange luminous aura that draws the eye” that is “easily visible from one hundred feet away with its serene, sinister, angelic radiance.” [4] The cap is smooth and usually described as viscid or tacky when wet.  This is to distinguish it from most of the other species in the Amanita genus that have warty patches on the cap from the dried out and cracking universal veil like the white dot warts on the bright red cap of the iconic fly agaric (Amanita muscaria).  The glowing purity of the whiteness is a reliable feature for initial field identification. Confirmation by looking for a picture or drawing of a white mushroom with a volva and annular stem ring using a field guide is another matter. One provides only Amanita verna or fool’s mushroom, prevalent only in spring (vernus in Latin). The common name implies that it fools the observer with its deception. [5] A second field guide provides both A. verna as the spring destroying angel, and Amanita virosa (virosus is poisonous in Latin) for mushrooms that appear in the fall with only a passing reference to A. bisporigera. [6] DNA sequencing of fungi has had a profound impact on the eighteenth-century Linnaean system basing taxonomy on physical similarity. It has been shown that all destroying angels of North America are A. bisporigera (with one additional species A. ocreata in California) and that A. verna and A. virosa are only found in Eurasia. Destroying angel is a universal common name for all species for the white mushrooms with volva.

The destroying angel is one of the deadliest mushrooms known. According to one account “misused as a cooking ingredient, its alabaster flesh has wiped out whole families.” [7] The toxic chemicals are called amatoxins (from the generic name Amanita), which are protein molecules made up of eight amino acids in a ring called a cyclopeptide with a molecular weight of about 900. The death dealing amatoxin variant is alpha-amanitin, which destroys RNA polymerase, a crucial metabolic enzyme. RNA polymerase transcribes the DNA blueprint, creating  messenger RNA that transport the codon amino acid recipe used  to make proteins on which all life depends. The ultimate result is rapid cell death. The gastrointestinal mucosa cells of the stomach, the hepatocytes of the liver, and the renal tubular cells of the kidneys are the most severely affected cells because they have the highest turnover rate and are rapidly depleted. The liver is most at risk because the hepatocytes that absorb alpha-amanitins are excreted with the bile and then reabsorbed. The initial stages of amatoxin poisoning start about ten hours after ingestion; the gastrointestinal mucosa cells are the first to be affected resulting in forcible eviction (aka vomiting) of the intruding poisons.  There follows a period of several days of calm as the stomach cells recover somewhat before the storm of  hepatic and renal debilitation. The third and final stage can in severe cases lead to the crescendo of convulsions, coma and death. The lethal dose for 50 percent of the population or LD50 is used by toxicologists as a benchmark for relative virulence. The LD50 for alpha-amanitin is 0.1 mg/kg.  A 70 kg adult will have a 50-50 chance of survival with a dose of 7 milligrams, the amount of alpha-amanitin in one small destroying angel. [8]

The North American Mycological Association (NAMA) received a total of 126 reports of amatoxin poisoning over a period of thirty years, about four annually. The fatality rate has historically been on the order of thirty percent attributed to liver and/or kidney failure; this number has improved over the last several decades to about five percent due to a better understanding of amatoxin physiology effects combined with aggressive therapy. The basic tenet of the treatment is to reduce the toxic concentration in the blood serum as rapidly as possible. Gastric lavation is used if the ingestion was recent enough followed by a thorough purging using emetics to induce vomiting and cathartics to induce evacuation of the bowels (essentially the same effect on the gastrointestinal mucosa cells to expel the poison).  Perhaps the most important therapy is the use of activated charcoal, as amatoxins have a high affinity for adsorption on its surface. Although there is no proven antidote, intravenous injections of penicillin have been used with some apparent benefit. A French physician named Bastien developed a three part procedure using intravenous injections of vitamin C and two types of  antibacterial drugs supplemented with penicillin to successfully treat 15 cases. To unequivocally prove its efficacy, he conducted the ultimate experiment by eating 70 grams of Amanita phalloides, the death cap cousin of the destroying angel and using the protocol on himself. [9] The most promising new treatment is silibinin, an extract of the blessed milk thistle (Silybum marianum), which is sold commercially as Legalon by a German pharmaceutical company. Liver transplant was once considered the last resort for amatoxin poisoning, but that may no longer be necessary. [10]

The destroying angel is not the only mushroom that produces amatoxin, nor is amatoxin the only substance produced by fungi that is inimical to humans. The identification of fungal toxins and the characterization of their imputed symptoms are among the most empirical of forensic science. The facts are based almost entirely on the anecdote. The identification of the mushroom that caused the condition under evaluation is usually a matter of conjecture since the victim has eaten the evidence. To add to the confusion, the alleged offending mushroom may have been consumed with a mixture of other wild foods and fungi gathered over a wide area in obscure nooks.  The dearth of fungal knowledge in the medical community contributes to uncertainly. Poison Control Centers (PCC) were established after World War II to deal with the proliferation of chemicals as clearing houses for information about poisons and their antidotes and treatment protocols. [11] Over the ensuing years, mushroom poisonings accounted for only one half of one percent of all PCC reports (1 in 200). Of those reported, only 10 percent included any information about the mushroom. Based on limited data, NAMA established a toxicology committee in 1985 and began to supplement the PCC data with a separate data base using the input from experienced mycologists and mushroom aficionados. The result to date is a more comprehensive accounting with fairly reliable identification of 80 percent of the mushrooms involved in poisoning. [12] This is a good start but has done little to assuage the beliefs of the general public that most if not all mushrooms are toadstools and that eating wild mushrooms is a fool’s errand, sometimes literally.

One example suffices to point out the irrational fear of amanita mushroom poisoning and the broader category of mycophobia. In 1991, the venerable French reference Petit Larousse Encyclopédie was recalled because the deadly amanita article lacked the appropriate symbol for poison. But this was not enough, since almost 200,000 copies had already been sold.  Several hundred students were hired to visit 6,000 stores throughout Europe and Canada to affix stickers with the appropriate symbol for poison on the pages and append a notice on the cover of the book that it was a new edition. [13]  History has impugned the mushroom as the source of the poison that has dispatched any number of notables, among them Claudius, the fourth Roman Emperor. The perpetrator is alleged to have been his fourth wife Agrippina who wanted her son Nero to succeed to the throne. The death is recounted by the philosopher Seneca the Younger in December 54 CE, only two months after the event occurred. According to his account, it happened quite quickly, the onset of illness and death being separated only by about an hour. [14] The mushroom assassination of Claudius is almost certainly apocryphal, as deadly mushrooms are relatively slow to act; those that act rapidly generally cause gastrointestinal distress that is rarely fatal. Hyperbole is not out of the question. One recent account attributes the disappearance of the Lost Colony of Roanoke to the relocation of the starving colonists to the island of Croatoan. Gorging themselves on the mushroom bounty that they found there, they died a horrible death of grotesque contortions. [15]

References:

1. McIlvaine, C. One Thousand American Fungi, Dover Publications, New York, 1973 pp 2-5

2. Roody. W. Mushrooms of West Virginia and the Central Appalachians, The University Press of Kentucky, Lexington, Kentucky, 2003, pp 62-63.

3. Lincoff, G. National Audubon Society Field Guide to North American Mushrooms, Alfred A. Knopf, New York, 1981. pp 551-552.

4. Russel, B. Field Guide to Wild Mushrooms of Pennsylvania and the Mid-Atlantic, The Penn State University Press, University Park, Pennsylvania, 1935, pp 67-69.

5. McKnight, K and McKnight, V.  Peterson Field Guide to Mushrooms of North America, Houghton Mifflin Company, Boston, 1987, pp 238-239, Plate 27.

6. Pacioni, G. (Lincoff, G, US editor) Guide to Mushrooms, Simon and Schuster, New York, 1981, pp 76-77.

7. Money, N. Mr. Bloomfield’s Orchard, Oxford University Press, Oxford. 2002 p 151

8. Hallen, H. et al. “Gene family encoding the major toxins of lethal Amanita mushrooms”. Proceedings of the National Academy of Sciences. 27 November 2007 Volume  104  Number 48  pp 19097–19101

9. Kendrick, B. The Fifth Kingdom, Focus Publishing, Newburyport, Massachusetts, 2000, pp 319-321.

10. Beug, M. in Fungi Magazine Volume 1 Number 2 Spring 2008. Beug is a Professor Emeritus at Evergreen State College and a member of the NAMA toxicology committee.

11. Wyckoff, A. “AAP Had First Hand in Poison Control Center” AAP News Sept. 2013 http://www.aappublications.org/content/34/10/45

12. Beug, M, et al “Thirty-Plus Years of Mushroom Poisoning: Summary of the Approximately 2,000 Reports in the NAMA Case Registry” Mcllvanea Volume 16 number 2 Fall 2006 pp 47-68.

13, Schaechter, E. In the Company of Mushrooms,  Harvard University Press, Cambridge, Massachusetts, 1997, pp 210-211.

14. Marmon, V. and Wiedemann, T. “The Death of Claudius” Journal of the Royal Society of Medicine, Volume 95, May 2002 pp. 260-261.

15. Spenser, S. “The First Case of Mass Mushroom Poisoning in the New World” Fungi Magazine, Volume 11, Number 4, Fall 2018, pp 30-33.

Inky Cap – Coprinoid mushrooms

Coprinus comatus shaggy mane Pyrenees 1509110
The Shaggy Mane is the most well known of the inky caps, and one of the few that remains in the genus Coprinus

Common Name:  Shaggy Mane, lawyer’s  wig, inky cap  – The unusual bullet shape of the cap bears some resemblance to the pate so that the cascading scales become disheveled locks or shaggy mane.

Scientific Name: Coprinus comatus – The generic name is from the Greek koprinos, meaning “of dung,” as many of its constituent fungi grow on animal feces. Comatus is Latin for “hairy,” referring to the texture of the cap. Family: Coprinaceae (now Agaricaceae)

Potpourri:  The Coprinaceae was widely known as the inky cap family for the notable and unique characteristic behavior of some of the larger species like the shaggy mane. Rather than open out into the umbrella shape of a typical mushroom for the air-borne dispersal of spores, the inky caps slowly dissolve into a black, gelatinous fluid that oozes slowly to the ground below. The gradual decomposition of  plant and mushroom tissues into a gooey mass is called deliquescence, a specialized case of decomposition. This term is sometimes applied to the inky caps but it is somewhat of a misnomer. Stinkhorns also prematurely degrade to a syrupy liquid that is redolent ― the aroma attracts insects that crawl through the muck and then fly away, dispersing the spores as they go.   The inky cap or  coprinoid mushrooms have a similar purpose absent the smell and the flies. The cap dissolves from the bottom up so that the gills and their attached spores are sequentially uncovered to allow for gradual spore dispersal by air currents. It is considered likely that this is an improvement over the typical mushroom arrangement with spore-bearing gills that extend from the underside of the cap. Once the caps open, the spores, which had been protected during the extension from their hypogeal origination in the mycelium, become exposed to environmental degradation.  The inky cap spores remain covered and thus protected until they are ready for deployment, probably as an evolutionary enhancement to improve survivability. [1]

It is wholly logical  that a peculiar characteristic like cap dissolution into inky black fluid would unite a group of fungi in having a common ancestor. The taxonomic system devised by Carolinus Linnaeus in the eighteenth century relied on such similarities to establish the hierarchical relationships that are still used today; the common traits defined and established family trees with genera of species  below and classes of orders above. Darwin’s observations of these similarities led to his evolutionary theory based on random mutations and survival of the fittest to explain trait radiation from an originator.  All was well with biology until about fifty years ago when the secret life of the genome was slowly but inexorably exposed. As is now well known and institutionalized, biological history is recorded in the arrangement of just four nucleotide bases: adenine A, cytosine C, guanine G, and thymine T (replaced by uracil U when transcribed by RNA). While the complexity of inky cap relationships is far from settled, the inky cap family Coprinaceae is no more. The shaggy mane, C. comatus, which had been the type specimen for the genus Coprinus, is one of the few that remain but it is now in a different family – Agarciaceae. Coprinus is therefore now the inky cap genus in a different family.

Coprinus quadrifidus 3 Scaly Inky Cao Columbia 200529
Scaly inky caps grow in clusters in hardwood debris and mulched areas

The coprinoid mushrooms that have a different DNA profile have been moved to three new genera named Coprinopsis, Coprinellus, and Parasola,  the first two retaining the scatological association of the original. Based on their DNA similarity to the genus Psathyrella they have been included in a newly created family named Psathyrellaceae. [2] The most common example of the newly minted “Coprinus-like” genus is Coprinopsis variegatus (also quadrifidus) , which fortunately goes by the much more mnemonic common name “scaly inky cap.” They appear in large clusters the day after a good soaking rain from decaying hardwood debris, the shaggy domed shapes like a conclave of bewigged nobility or a battery of ballistic missiles. Although they are considered edible, this would only be for the adventuresome, as they are noted for frequently having an unpleasant odor and taste presumably extracted from the substrate debris from which they erupted.

Coprinus micaceous Mica Cap gills AT Rt7 180919
The mica cap has veil fragments on the cap that glisten like mica.

The “little coprinuses” of the new genus Coprinellus are at the opposite end of the larger, maned sporocarps of Coprinus and Coprinus-like fungi of the once majestic inky cap family dining room table. [3] While retaining the deliquescing inkiness of spore dispersal, the bullet shape is absent in most species in favor of a more typical cap and stem mushroom arrangement. The most well known is the mica cap, C. micaceus, which takes its name from the Latin micare meaning “to flash or sparkle.” The silicate mineral mica is a primary constituent of igneous rocks like granite that are characterized by scintillation. Mica caps have evanescent speckles that are the remnants of the partial veil, a barrier employed by some mushrooms to protect the spore-bearing gills on the underside of the cap until the stem is fully extended and the cap opens. They are widely dispersed and quite common on stumps and woody debris and even indoors … David Aurora noted that “bountiful crops sprout periodically from the woodwork of a popular café in Santa Cruz, California.” [4] C. micaceus may be the world’s first scientifically described mushroom, appearing as a woodcut illustration in Rariorum plantarum historia by Carolus Clusius that was published in the early seventeenth century. [5]

Even as it has been removed from the patriarchal position as the quintessence of the inky cap family that is no more, the shaggy mane Coprinus comatus retains it notoriety. It is certainly the inspiration for a stanza on mushrooms in Shelley’s The Sensitive Plant:

 

Their moss rotted off them, flake by flake

Till the thick stalk stuck like a murderer’s stake

Where rags of loose flesh yet tremble on high’

Infecting the winds that wander by [6]

 

The importance of inky caps extends to the laboratory and they have accordingly been the subject of numerous foundational fungal research efforts over the years. particularly in the area of sporulation. The formation of the “ink” is due to gill autolysis, the removal of interference for sequential spore ejection in all gilled mushrooms. The spores are held in place by club-shaped structures called basidia that extend outward on both faces of the vertical gill. The inky caps are masters of spores and spore release ― the shaggy mane produces roughly nine thousand million spores in about three days. That works out to 30,000 spores per second.[7]

Dung is a potent source of nutrition for significant swaths of the Kingdom Fungi, the genus name Coprinus is not coincidental. Although feces as food is repugnant to people, coprophagy is standard fare for some fungi, some beetles … one of the scarabs is named dung beetle… and rabbits. For one thing there is a lot of it, deposited daily by roving bands of herbivores leaving cow pies and horse “road apples” in their wake. While most of the proteins are gone, it is replete with cellulose that animals can’t digest but fungi can and nitrogen, a vital element for all things living. Dung can contain up to four percent nitrogen, which is more than the original ingested plant material. There are almost two hundred genera of fungi, mostly ascomycetes or cup fungi, that are primarily coprophagous. One of the more intriguing examples of nature’s insidious exploitation is the zygomycete Pilobolus crystallinus.  Called “the hat thrower,” it is one of several species of fungi that have evolved to grow in dung, shoot their spores up to two meters away toward light, and germinate in the grass away from the dung pile where they are consumed by grazing animals. Passing unaffected through the animal’s digestive system, the spores are deposited in new dung in a new place to perpetuate the cycle. Once the zygomycetes and ascomycetes are done, basidiomycetes like the coprinoid mushrooms take over, a succession based on resources needed for fruiting body formation. [8] Ultimately, the dung is recycled ecologically ― a good thing for otherwise we would be buried in it.

Shaggy manes are one of the more noted edible mushrooms, characterized as “choice” in the more popular mushroom field guides, one of which calls it “an excellent substitute for asparagus, it can also be pickled.” [9] As with all edible fungi, however,  caveats apply and there are  doppelgängers that entice the ill-informed neophyte. In this case, it is the “alcohol inky,” Coprinopsis (nee Coprinus)  atramentarius that looks more or less like a shaggy mane with a haircut. As mushrooms are somewhat variable in appearance according to age and disposition, one might well mistake this for its cousin C. comatus. This would not be a serious problem since it is also edible unless your gourmet meal of wild mushrooms includes a glass of wine. The problem is that the alcohol inky contains the toxin coprine, which is similar in effect to disulfiram, the chemical used  in drugs administered to enforce alcohol abstinence in those suffering from its addiction. [10] When alcohol is consumed under normal conditions by most people, it is converted to acetaldehyde which is then metabolized to acetate resulting in a pleasant light-headed feeling of mild euphoria. Coprine blocks the metabolic pathway so that acetaldehyde builds up in the body causing “flushing of the face, headache, nausea, vomiting, chest pain, weakness, blurred vision, mental confusion, sweating, choking, breathing difficulty, and anxiety”  known as the disulfiram-ethanol reaction. [11] Eating wild fungi has always been a challenge, but some find it worth the time and trouble. Once the subtleties of identification have been mastered, it is the joy of the hunt that prevails.

References:

  1. Roody, W. Mushrooms of West Virginia and the Central Appalachians, The University of Kentucky Press, Lexington, Kentucky, 2003, pp 268-269
  2. Redhead S. et al. “Coprinus Pers. and the disposition of Coprinus species sensu lato”. Taxon. 1 February 2001 Volume 50 (1) pp 203–241
  3. Volk, T. Coprinus comatus, Fungus of the Month May 2004 available at http://botit.botany.wisc.edu/toms_fungi/may2004.html
  4. Aurora, D. Mushrooms Demystified, Ten Speed Press, Berkeley California, 1986. pp 348-349
  5. Bulliard J.. Herbier de la France [Guide to the Herbs of France] 1786. pp. 241–88, plate 246. See . https://www.biodiversitylibrary.org/item/25508#page/16/mode/1up
  6. Money, N. Mr. Bloomfield’s Orchard, Oxford University Press, New York, 2002. p 17.
  7. Carlile, M., Watkinson, S. and Gooday, G. The Fungi, 2nd edition, Elsevier Academic Press, London, 2006, pp 61-63.
  8. Kendrick, B. The Fifth Kingdom, 3rd edition, Focus Publishing, Newburyport, Massachusetts, 2000. pp 33-34, 184-185.
  9. Lincoff, G. National Audubon Society Field Guide to North American Mushrooms, Alfred A. Knopf, New York, 1981, 99 596-598.
  10. Money, op cit. p 157.
  11. https://medlineplus.gov/druginfo/meds/a682602.html