Amanita phalloides

Amanita phalloides (/æməˈnaɪtə

fəˈlɔɪdiːz/), commonly known as the
death cap, is a deadly poisonous
basidiomycete fungus, one of many in
the genus Amanita. Widely distributed
across Europe, but now sprouting in
other parts of the world, A. phalloides
forms ectomycorrhizas with various
broadleaved trees. In some cases, the
death cap has been introduced to new
regions with the cultivation of non-native
species of oak, chestnut, and pine. The
large fruiting bodies (mushrooms)
appear in summer and autumn; the caps
are generally greenish in colour with a
white stipe and gills. The cap colour is
variable, including white forms, and is
thus not a reliable identifier.
 Amanita phalloides

In Piacenza, Italy

Scientific classification

Kingdom: Fungi

Division: Basidiomycota

Class: Agaricomycetes

Order: Agaricales

Family: Amanitaceae

Genus: Amanita
Species: A. phalloides
Binomial name

Amanita phalloides

(Vaill. ex Fr.) Link (1833)

Amanita phalloides
Mycological characteristics
gills on hymenium

cap is convex
or flat

hymenium is free

stipe has a ring and

volva

spore print is white

ecology is
mycorrhizal

edibility: deadly
These toxic mushrooms resemble
several edible species (most notably
Caesar's mushroom and the straw
mushroom) commonly consumed by
humans, increasing the risk of accidental
poisoning. Amatoxins, the class of toxins
found in these mushrooms, are
thermostable: they resist changes due to
heat, so their toxic effects are not
reduced by cooking.

A. phalloides is one of the most

poisonous of all known mushrooms. It is
estimated that as little as half a
mushroom contains enough toxin to kill
an adult human. It has been involved in
the majority of human deaths from
mushroom poisoning,[1] possibly
including Roman Emperor Claudius in AD
54 and Holy Roman Emperor Charles VI
in 1740. It has been the subject of much
research and many of its biologically
active agents have been isolated. The
principal toxic constituent is α-Amanitin,
which causes liver and kidney failure.

Taxonomy
The death cap is named in Latin as such
in the correspondence between the
English physician Thomas Browne and
Christopher Merrett.[2] Also, it was
described by French botanist Sébastien
Vaillant in 1727, who gave a succinct
phrase name "Fungus phalloides,
annulatus, sordide virescens, et patulus"—
a recognizable name for the fungus
today.[3] Though the scientific name
phalloides means "phallus-shaped", it is
unclear whether it is named for its
resemblance to a literal phallus or the
stinkhorn mushrooms Phallus.
In 1821,
Elias Magnus Fries described it as
Agaricus phalloides, but included all
white amanitas within its description.[4]
Finally, in 1833, Johann Heinrich Friedrich
Link settled on the name Amanita
phalloides,[5] after Persoon had named it
Amanita viridis 30 years earlier.[6][7]
Although Louis Secretan's use of the
name A. phalloides predates Link's, it has
been rejected for nomenclatural
purposes because Secretan's works did
not use binomial nomenclature
consistently;[8][9] some taxonomists have,
however, disagreed with this
opinion.[10][11]

Amanita phalloides is the type species of

Amanita section Phalloideae, a group
that contains all of the deadly poisonous
Amanita species thus far identified. Most
notable of these are the species known
as destroying angels, namely A. virosa,
A. bisporigera and A. ocreata, as well as
the fool's mushroom (A. verna). The term
"destroying angel" has been applied to
A. phalloides at times, but "death cap" is
by far the most common vernacular
name used in English. Other common
names also listed include "stinking
amanita"[12] and "deadly amanita".[13]

A rarely appearing, all-white form was

initially described A. phalloides f. alba by
Max Britzelmayr,[14][15] though its status
has been unclear. It is often found
growing amid normally colored death
caps. It has been described, in 2004, as a
distinct variety and includes what was
termed A. verna var. tarda.[16] The true
A. verna fruits in spring and turns yellow
with KOH solution, whereas A. phalloides
never does.[17]
Description
The death cap has a large and imposing
epigeous (aboveground) fruiting body
(basidiocarp), usually with a pileus (cap)
from 5 to 15 centimetres (2 to 57⁄8
inches) across, initially rounded and
hemispherical, but flattening with age.[18]
The color of the cap can be pale-green,
yellowish-green, olive-green, bronze, or
(in one form) white; it is often paler
toward the margins, which can have
darker streaks;[19] it is also often paler
after rain. The cap surface is sticky when
wet and easily peeled, a troublesome
feature, as that is allegedly a feature of
edible fungi.[20] The remains of the partial
veil are seen as a skirtlike, floppy annulus
usually about 1 to 1.5 cm (3⁄8 to 5⁄8 in)
below the cap. The crowded white
lamellae (gills) are free. The stipe is
white with a scattering of grayish-olive
scales and is 8 to 15 cm (31⁄8 to 57⁄8 in)
long and 1 to 2 cm (3⁄8 to 3⁄4 in) thick,
with a swollen, ragged, sac-like white
volva (base).[18] As the volva, which may
be hidden by leaf litter, is a distinctive
and diagnostic feature, it is important to
remove some debris to check for it.[21]

The smell has been described as initially

faint and honey-sweet, but strengthening
over time to become overpowering,
sickly-sweet and objectionable.[22] Young
specimens first emerge from the ground
resembling a white egg covered by a
universal veil, which then breaks, leaving
the volva as a remnant. The spore print is
white, a common feature of Amanita. The
transparent spores are globular to egg-
shaped, measure 8–10 μm (0.3–0.4 mil)
long, and stain blue with iodine.[22] The
gills, in contrast, stain pallid lilac or pink
with concentrated sulfuric acid.[23][24]

Biochemistry
α-Amanitin

β-Amanitin, where an amide of the α-Amanitin has been replaced by a carboxylic acid (lower left corner)

The species is now known to contain two

main groups of toxins, both multicyclic
(ring-shaped) peptides, spread
throughout the mushroom tissue: the
amatoxins and the phallotoxins. Another
toxin is phallolysin, which has shown
some hemolytic (red blood cell–
destroying) activity in vitro. An unrelated
compound, antamanide, has also been
isolated.
Amatoxins consist of at least eight
compounds with a similar structure, that
of eight amino-acid rings; they were
isolated in 1941 by Heinrich O. Wieland
and Rudolf Hallermayer of the University
of Munich.[25] Of the amatoxins, α-
Amanitin is the chief component and
along with β-amanitin is likely
responsible for the toxic effects.[26][27]
Their major toxic mechanism is the
inhibition of RNA polymerase II, a vital
enzyme in the synthesis of messenger
RNA (mRNA), microRNA, and small
nuclear RNA (snRNA). Without mRNA,
essential protein synthesis and hence
cell metabolism grind to a halt and the
cell dies.[28] The liver is the principal
organ affected, as it is the organ which is
first encountered after absorption in the
gastrointestinal tract, though other
organs, especially the kidneys, are
susceptible.[29] The RNA polymerase of
Amanita phalloides is insensitive to the
effects of amatoxins, so the mushroom
does not poison itself.[30]

The phallotoxins consist of at least seven

compounds, all of which have seven
similar peptide rings. Phalloidin was
isolated in 1937 by Feodor Lynen,
Heinrich Wieland's student and son-in-
law, and Ulrich Wieland of the University
of Munich. Though phallotoxins are
highly toxic to liver cells,[31] they have
since been found to add little to the
death cap's toxicity, as they are not
absorbed through the gut.[28]
Furthermore, phalloidin is also found in
the edible (and sought-after) blusher
(A. rubescens).[25] Another group of
minor active peptides are the virotoxins,
which consist of six similar monocyclic
heptapeptides.[32] Like the phallotoxins,
they do not induce any acute toxicity
after ingestion in humans.[28]

The genome of the death cap has been

sequenced.[33]
Death caps in French deciduous wood

A young death cap emerging from its

universal veil
 Young "death cap" mushrooms in Poland,
with matchbox for scale

Similarity to edible species

A. phalloides is similar to the edible

paddy straw mushroom (Volvariella
volvacea)[34] and A. princeps, commonly
known as "white Caesar".[35]

Some may mistake juvenile death caps

for edible puffballs[36][37] or mature
specimens for other edible Amanita
species, such as A. lanei, so some
authorities recommend avoiding the
collecting of Amanita species for the
table altogether.[38] The white form of
A. phalloides may be mistaken for edible
species of Agaricus, especially the young
fruitbodies whose unexpanded caps
conceal the telltale white gills; all mature
species of Agaricus have dark-colored
gills.[39]

In Europe, other similarly green-capped

species collected by mushroom hunters
include various green-hued brittlegills of
the genus Russula and the formerly
popular Tricholoma equestre, now
regarded as hazardous owing to a series
of restaurant poisonings in France.
Brittlegills, such as Russula heterophylla,
R. aeruginea, and R. virescens, can be
distinguished by their brittle flesh and the
lack of both volva and ring.[40] Other
similar species include A. subjunquillea in
eastern Asia and A. arocheae, which
ranges from Andean Colombia north at
least as far as central Mexico, both of
which are also poisonous.

Distribution and habitat

The death cap is native to Europe, where
it is widespread.[41] It is found from the
southern coastal regions of Scandinavia
in the north, to Ireland in the west, east to
Poland and western Russia,[16] and south
throughout the Balkans, in Greece, Italy,
Spain, and Portugal in the Mediterranean
basin, and in Morocco and Algeria in
north Africa.[42] In west Asia, it has been
reported from forests of northern Iran.[43]
There are records from further east in
Asia but these have yet to be confirmed
as A. phalloides.[44]

By the end of the 19th century, Charles

Horton Peck had reported A. phalloides in
North America.[45] In 1918, samples from
the eastern United States were identified
as being a distinct though similar
species, A. brunnescens, by George
Francis Atkinson of Cornell University.[25]
By the 1970s, it had become clear that
A. phalloides does occur in the United
States, apparently having been
introduced from Europe alongside
chestnuts, with populations on the West
and East Coasts.[25][46] A 2006 historical
review concluded the East Coast
populations were inadvertently
introduced, likely on the roots of other
purposely imported plants such as
chestnuts.[47] The origins of the West
Coast populations remained unclear, due
to scant historical records,[44] but a 2009
genetic study provided strong evidence
for the introduced status of the fungus
on the west coast of North America.[48]
Observations of various collections of
A. phalloides, from conifers rather than
native forests, have led to the hypothesis
that the species was introduced to North
America multiple times. It is
hypothesized that the various
introductions led to multiple genotypes
which are adapted to either oaks or
conifers.[49]

A. phalloides has been conveyed to new

countries across the Southern
Hemisphere with the importation of
hardwoods and conifers. Introduced
oaks appear to have been the vector to
Australia and South America; populations
under oaks have been recorded from
Melbourne and Canberra[50][51][52] (where
two people died in January 2012, of four
who were poisoned)[53] and Adelaide,[54]
as well as Uruguay.[55] It has been
recorded under other introduced trees in
Argentina[56]Pine plantations are
associated with the fungus in
Tanzania[57] and South Africa, where it is
also found under oaks and poplars and
ChileValenzuella, E.; Moreno, G.; Jeria, M.
(1992). "Amanita phalloides en bosques
de Pinus radiata de la IX Region de Chile:
taxonomia, toxinas, metodos de
dedection, intoxicacion faloidiana".
Boletín Micológico. 7: 17–21. ISSN 0716-
114X (https://www.worldcat.org/issn/07
16-114X) .[58] A number of deaths in
India have been attributed to it.[59]
Ecology
It is ectomycorrhizally associated with
several tree species and is symbiotic
with them. In Europe, these include
hardwood and, less frequently, conifer
species. It appears most commonly
under oaks, but also under beeches,
chestnuts, horse-chestnuts, birches,
filberts, hornbeams, pines, and
spruces.[14] In other areas, A. phalloides
may also be associated with these trees
or with only some species and not
others. In coastal California, for example,
A. phalloides is associated with coast live
oak, but not with the various coastal pine
species, such as Monterey pine.[60] In
countries where it has been introduced, it
has been restricted to those exotic trees
with which it would associate in its
natural range. There is, however,
evidence of A. phalloides associating
with hemlock and with genera of the
Myrtaceae: Eucalyptus in Tanzania[57]
and Algeria,[42] and Leptospermum and
Kunzea in New Zealand,[14][61] suggesting
that the species may have invasive
potential.[44] It may have also been
anthropogenically introduced to the
island of Cyprus, where it has been
documented to fruit within Corylus
avellana plantations.[62]

Toxicity
Warning sign in Canberra, Australia

As the common name suggests, the

fungus is highly toxic, and is responsible
for the majority of fatal mushroom
poisonings worldwide.[1][63] Its
biochemistry has been researched
intensively for decades,[25] and 30 grams
(1.1 ounces), or half a cap, of this
mushroom is estimated to be enough to
kill a human.[64] On average, one person
dies a year in North America from death
cap ingestion.[35] The toxins of the death
cap mushrooms primarily target the liver,
but other organs, such as the kidneys, are
also affected. Symptoms of death cap
mushroom toxicity usually occur 6 to 12
hours after ingestion.[65] Symptoms of
ingestion of the death cap mushroom
may include nausea and vomiting, which
is then followed by jaundice, seizures,
and coma which will lead to death. The
mortality rate of ingestion of the death
cap mushroom is believed to be around
10–30%.[66]

Some authorities strongly advise against

putting suspected death caps in the
same basket with fungi collected for the
table and to avoid even touching
them.[20][67] Furthermore, the toxicity is
not reduced by cooking, freezing, or
drying.[68]

Poisoning incidents usually result from

errors in identification. Recent cases
highlight the issue of the similarity of
A. phalloides to the edible paddy straw
mushroom (Volvariella volvacea), with
East- and Southeast-Asian immigrants in
Australia and the West Coast of the U.S.
falling victim. In an episode in Oregon,
four members of a Korean family
required liver transplants.[34] Many North
American incidents of death cap
poisoning have occurred among Laotian
and Hmong immigrants, since it is easily
confused with A. princeps ("white
Caesar"), a popular mushroom in their
native countries.[35] Of the 9 people
poisoned in the Canberra region between
1988 and 2011, three were from Laos
and two were from China.[68] In January
2012, four people were accidentally
poisoned when death caps (reportedly
misidentified as straw fungi, which are
popular in Chinese and other Asian
dishes) were served for dinner in
Canberra; all the victims required hospital
treatment and two of them died, with a
third requiring a liver transplant.[69]
Signs and symptoms

Death caps have been reported to taste

pleasant.[25][70] This, coupled with the
delay in the appearance of symptoms—
during which time internal organs are
being severely, sometimes irreparably,
damaged—makes it particularly
dangerous. Initially, symptoms are
gastrointestinal in nature and include
colicky abdominal pain, with watery
diarrhea, nausea, and vomiting, which
may lead to dehydration if left untreated,
and, in severe cases, hypotension,
tachycardia, hypoglycemia, and acid–
base disturbances.[71][72] These first
symptoms resolve two to three days
after the ingestion. A more serious
deterioration signifying liver involvement
may then occur—jaundice, diarrhea,
delirium, seizures, and coma due to
fulminant liver failure and attendant
hepatic encephalopathy caused by the
accumulation of normally liver-removed
substance in the blood.[12] Kidney failure
(either secondary to severe
hepatitis[32][73] or caused by direct toxic
kidney damage[28]) and coagulopathy
may appear during this stage. Life-
threatening complications include
increased intracranial pressure,
intracranial bleeding, pancreatic
inflammation, acute kidney failure, and
cardiac arrest.[71][72] Death generally
occurs six to sixteen days after the
poisoning.[74]

Mushroom poisoning is more common in

Europe than in North America.[75] Up to
the mid-20th century, the mortality rate
was around 60–70%, but this has been
greatly reduced with advances in medical
care. A review of death cap poisoning
throughout Europe from 1971 to 1980
found the overall mortality rate to be
22.4% (51.3% in children under ten and
16.5% in those older than ten).[76] This
has fallen further in more recent surveys
to around 10–15%.[77]

Treatment
Consumption of the death cap is a
medical emergency requiring
hospitalization. The four main categories
of therapy for poisoning are preliminary
medical care, supportive measures,
specific treatments, and liver
transplantation.[78]

Preliminary care consists of gastric

decontamination with either activated
carbon or gastric lavage; due to the delay
between ingestion and the first
symptoms of poisoning, it is common for
patients to arrive for treatment many
hours after ingestion, potentially reducing
the efficacy of these interventions.[78][79]
Supportive measures are directed
towards treating the dehydration which
results from fluid loss during the
gastrointestinal phase of intoxication and
correction of metabolic acidosis,
hypoglycemia, electrolyte imbalances,
and impaired coagulation.[78]

No definitive antidote is available, but

some specific treatments have been
shown to improve survivability. High-
dose continuous intravenous penicillin G
has been reported to be of benefit,
though the exact mechanism is
unknown,[76] and trials with
cephalosporins show promise.[80][81]
Some evidence indicates intravenous
silibinin, an extract from the blessed milk
thistle (Silybum marianum), may be
beneficial in reducing the effects of death
cap poisoning. A long-term clinical trial
of intravenous silibinin began in the US in
2010.[82] Silibinin prevents the uptake of
amatoxins by liver cells, thereby
protecting undamaged liver tissue; it also
stimulates DNA-dependent RNA
polymerases, leading to an increase in
RNA synthesis.[83][84][85] According to one
report[86] based on a treatment of 60
patients with silibinin, patients who
started the drug within 96 hours of
ingesting the mushroom and who still
had intact kidney function all survived. As
of February 2014 supporting research
has not yet been published.
SLCO1B3 has been identified as the
human hepatic uptake transporter for
amatoxins; moreover, substrates and
inhibitors of that protein—among others
rifampicin, penicillin, silibinin,
antamanide, paclitaxel, ciclosporin and
prednisolone—may be useful for the
treatment of human amatoxin
poisoning.[87]

N-Acetylcysteine has shown promise in

combination with other therapies.[88]
Animal studies indicate the amatoxins
deplete hepatic glutathione;[89] N-
acetylcysteine serves as a glutathione
precursor and may therefore prevent
reduced glutathione levels and
subsequent liver damage.[90] None of the
antidotes used have undergone
prospective, randomized clinical trials,
and only anecdotal support is available.
Silibinin and N-acetylcysteine appear to
be the therapies with the most potential
benefit.[78] Repeated doses of activated
carbon may be helpful by absorbing any
toxins returned to the gastrointestinal
tract following enterohepatic
circulation.[91] Other methods of
enhancing the elimination of the toxins
have been trialed; techniques such as
hemodialysis,[92] hemoperfusion,[93]
plasmapheresis,[94] and peritoneal
dialysis[95] have occasionally yielded
success, but overall do not appear to
improve outcome.[28]

In patients developing liver failure, a liver

transplant is often the only option to
prevent death. Liver transplants have
become a well-established option in
amatoxin poisoning.[71][72][96] This is a
complicated issue, however, as
transplants themselves may have
significant complications and mortality;
patients require long-term
immunosuppression to maintain the
transplant.[78] That being the case, the
criteria have been reassessed, such as
onset of symptoms, prothrombin time
(PT), serum bilirubin, and presence of
encephalopathy, for determining at what
point a transplant becomes necessary
for survival.[97][98][99] Evidence suggests,
although survival rates have improved
with modern medical treatment, in
patients with moderate to severe
poisoning, up to half of those who did
recover suffered permanent liver
damage.[100] A follow-up study has
shown most survivors recover
completely without any sequelae if
treated within 36 hours of mushroom
ingestion.[101]

Notable victims
 Ce plat de champignons a
changé la destinée de l'Europe.

[This dish of mushrooms

changed the destiny of Europe.]

— Voltaire, Mémoires

Several historical figures may have died

from A. phalloides poisoning (or other
similar, toxic Amanita species). These
were either accidental poisonings or
assassination plots. Alleged victims of
this kind of poisoning include Roman
Emperor Claudius, Pope Clement VII, the
Russian tsaritsa Natalia Naryshkina, and
Holy Roman Emperor Charles VI.[102]
R. Gordon Wasson recounted[102] the
details of these deaths, noting the
likelihood of Amanita poisoning. In the
case of Clement VII, the illness that led to
his death lasted five months, making the
case inconsistent with amatoxin
poisoning. Natalya Naryshkina is said to
have consumed a large quantity of
pickled mushrooms prior to her death. It
is unclear whether the mushrooms
themselves were poisonous or if she
succumbed to food poisoning.

Charles VI experienced indigestion after

eating a dish of sautéed mushrooms.
This led to an illness from which he died
10 days later—symptomatology
consistent with amatoxin poisoning. His
death led to the War of the Austrian
Succession. Noted Voltaire, "this dish of
mushrooms changed the destiny of
Europe."[102][103]

The case of Claudius's poisoning is more

complex. Claudius was known to have
been very fond of eating Caesar's
mushroom. Following his death, many
sources have attributed it to his being fed
a meal of death caps instead of Caesar's
mushrooms. Ancient authors, such as
Tacitus and Suetonius, are unanimous
about poison having been added to the
mushroom dish, rather than the dish
having been prepared from poisonous
mushrooms. Wasson speculated the
poison used to kill Claudius was derived
from death caps, with a fatal dose of an
unknown poison (possibly a variety of
nightshade) being administered later
during his illness.[102][104] Other historians
have speculated that Claudius may have
died of natural causes.

See also
Fungi
portal

List of Amanita species

List of deadly fungi

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Cited texts

Benjamin, Denis R. (1995).

Mushrooms: Poisons and Panaceas—A
Handbook for Naturalists, Mycologists
 and Physicians. New York: WH
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7167-2600-5.
Jordan, Peter; Wheeler, Steven (2001).
The Ultimate Mushroom Book. London:
Hermes House. ISBN 978-1-85967-
092-7.
Zeitlmayr, Linus (1976). Wild
Mushrooms: An Illustrated Handbook.
Hertfordshire: Garden City Press.
ISBN 978-0-584-10324-3.

External links
Wikimedia Commons has media
related to Amanita phalloides.

UK Telegraph Newspaper (September

2008) - One woman dead, another
critically ill after eating Death Cap
fungi (https://www.telegraph.co.uk/ne
ws/2978434/One-woman-dead-and-an
other-seriously-ill-after-eating-Death-Ca
p-mushrooms.html)
AmericanMushrooms.com - The Death
Cap Mushroom Amanita phalloides (htt
p://americanmushrooms.com/deathca
p.htm)
Amanita phalloides: the death cap (htt
p://www.mushroomexpert.com/amanit
a_phalloides.html)
Amanita phalloides: Invasion of the
Death Cap (http://www.bayareamushro
oms.org/mushroommonth/amanita_p
halloides.html)
Key to species of Amanita Section
Phalloideae from North and Central
America - Amanita studies website (htt
p://pluto.njcc.com/~ret/amanita/key.di
r/phallimb.html)
California Fungi—Amanita phalloides
(http://www.mykoweb.com/CAF/speci
es/Amanita_phalloides.html)
Death cap in Australia - ANBG website
(http://www.anbg.gov.au/fungi/deathc
ap.html)
On the Trail of the Death Cap
Mushroom (https://www.npr.org/templ
ates/story/story.php?storyId=725132
7) from National Public Radio
 "Amanita phalloides" (https://www.eol.
org/pages/1009706) at the
Encyclopedia of Life

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