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Introduction:
Welcome to the Amanita Studies site. This page and subordinate
pages provide data on species of the macrofungal, largely
ectomycorrhizal genus Amanita
(Amanitaceae, Agaricales, Basidiomycetes, Fungi) supplemented by
monotone or color illustrations of the fungi wherever possible. This
is likely always to be a work in progress. The "Site News" feature
(see the site navigation block on the left) can be used to learn of
recent additions, deletions, or other changes to the
site.
[ Note: The site is prepared to be viewed best
via Mozilla Firefox® with the browser window dimensions maximized
(press F11 in Firefox). Firefox can be downloaded without
charge.]
Three levels of detail are being implemented
for the data on taxa of Amanita:
- The first level of detail is a good quality
illustration.
- The second level of detail is such an
illustration (if one can be found) and a brief species description
(with restrained use of Latinate terms) and emphasizing characters
visible to the naked eye or with a 10X lens -- although
microscopic data (especially information about spores and basidial
clamps) is also included when available.
- The third level of detail includes both of
the above (when an illustration is available) as well as a full
technical description including illustrations of some microscopic
characters (e.g., see A.
umbrinolutea in Amanita sect.
Vaginatae).
At present, the section level lists
include 545 names of taxa. We have done
our best to exclude synonyms from section level lists, but there are
probably several cases of taxonomic synonyms still to be found in
those lists. We have achieved detail level "2" (some descriptions
needing further work) for 517 of the listed taxa.
The editors welcome offers of assistance in the
support and expansion of this website. Persons wishing to provide
technical information on, or illustrations of, additional taxa
should inquire by email
or by postal service to either of the editors. Illustrations will be
gratefully accepted; but, for the most part, can only be used on a
species page if they are accompanied by well-dried voucher material
of the pictured collection.
Key: ["t.b.d." = "to be
developed"]
Background tile depicts A. muscaria
subsp. flavivolvata of North and Central America.
Tiles in navigation column (at left) depict A. crocea of Europe.
Species
pages: The core of taxonomic information on this
site is presented on "species pages" as described in the
introduction, above. By clicking on the name
of a section in the following annotated list, the viewer of this
page will be transferred to an alphabetical list of all species
(and some infraspecific taxa) accepted by the editors in the
selected section. A few names are provisional, and a few are
in common usage despite being invalid. For information about
names misapplied in various geographic areas and for more
information concerning (at least) the number of unnamed taxa in
those areas, the viewer is directed to the "Keys & Checklists"
link in
the site navigating column (pale yellow) to the left. The Amanita Studies site's checklists/picture
books provide illustration-based access to the same species pages
that are accessible taxonomically here as well as by use of the
intrasite search function.
The family Amanitaceae R. Heim ex Pouzar is typified by the genus Amanita and presently comprises four
genera: Amanita,
Amarrendia Bougher & Lebel, Limacella Earle, and Torrendia Bres. Recent (including
some unpublished) molecular studies concur with the morphological
view that Limacella is a distinct
genus; however, Amarrendia and Torrendia seem rather likely to comprise
hypogeous and secotioid (respectively) species that evolved from
multiple ancestors in multiple sections of the genus Amanita.
The genus Amanita
is divided into two subgenera (well-supported both morphologically
and molecularly) and seven sections according to a conservative
viewing of modern taxonomy. In his publications, Yang has
made the segregation of Amanita sect.
Caesarea from sect. Vaginatae for a number of years.
The editors are now in agreement on this point, and the web site
has been changed to reflect a genus with seven
sections.
To continue with a taxonomic summary of the
Amanitaceae, scroll down or select a
taxonomic grouping from the following links/lists:
Amanita
is defined by the combination of (1) longitudinally
acrophysalidic stipe tissue [with inflated cells
elongate-clavate, terminal in most species (in terminal
chains in others)]; (2) divergent lamella trama; and (3)
schizohymenial development (unique to Amanita among all agarics).
Essentially, the "button" of an Amanita includes no empty spaces;
the stem (stipe), cap (pileus), gills (lamellae), and
veil(s) develop in place and the tissues dividing these
parts of the fruiting body become friable and/or die to
enable the unhindered expansion of the maturing mushroom.
The type
species of the genus is A.
muscaria (L.: Fr.) Lam. [ =Agaricus
muscarius L. (1753) ] in Amanita [subgenus Amanita] section Amanita.
[NB:
Images and well-documented dried collections of material
from outside their respective regional collecting areas are
sought by both editors.]
The seven
section names in common use are listed below under the
appropriate subgenus name:
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[ subgenus Amanita ]
[ section Amanita ] [ section Vaginatae ] [ section Caesareae ]
[
subgenus Lepidella ]
[ section Lepidella ] [ section Amidella ] [ section Phalloideae ] [ section Validae ]
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Amarrendia - The species of
this genus are hypogeous ("subterranean and truffle
like"). At present, the genus is known only from
Australia. The genus is largely based on shared
elements of gross morphology. Molecular work has
excluded several morphologically similar hypogeous entities
originally assigned to Amarrendia that did not belong in
the Amanitaceae. Detailed
morphological workups on Amarrendia material have not been
completed to our knowledge. Evidence suggests that the
truffle-like taxa in Amanita
have descended from an ancestor or ancestors assignable to
Amanita sect. Caesareae [ key (over 540 Kb PDF) ]. The
editors of these pages presently favor recombining all
amanitoid taxa of Amarrendia in
Amanita.
The type species of Amarrendia is A. oleosa Bougher & Lebel
(2002).
[NB: Images and well-documented
dried collections are sought by both editors.] |
  Limacella - Species of the
genus Limacella are strongly
differentiated from the genus Amanita by their mode of
development--the mode is not
schizohymenial. The lamellae of the taxa for which
investigation has been reported in the literature grow down
into empty space from the developing pileus. As a
result, unlike the species of Amanita, limacellas have a fertile
edge on their gills (lamellae). In Limacella, the analog of the
universal veil of Amanita is a
glutinous matrix supported by hyphae that arise NOT from a
pileipellis (i.e., cuticle or cap skin), but from a dense
layer in the uppermost part of the pileus context (i.e., cap
flesh). Indeed, as in most taxa of Amanita [sect. Lepidella] subsect. Vittadiniae there is no pileipellis
present in Limacella. A
membranous partial veil (i.e., ring, skirt, or annulus) is
present in some species.
The type species of Limacella is Agaricus delicatus Pers. : Fr.
(1821).
[NB: Images and well-documented
dried collections of Limacella
are sought by both editors.] |
 Torrendia - The species of
this genus are (1) secotioid; (2) expand from within a
membranous, universal veil; (3) have longitudinally
acrophysalidic stipe tissue (as in Amanita); (4) have inamyloid spores
(with one possible exception); and (5) have clamps on the
bases of their basidia (with the same possible exception).
Taxa of the Mediterranean region and Australia have been
assigned to Torrendia. At
least some of the taxa of this genus (including the type
species) appear to have had ancestors in common with species
of Amanita section Caesareae [ key (over 540 Kb PDF) ]. As
in the case of Amarrendia (above), the status of
the genus Torrendia is under
on-going investigation. At present, the editors of
these pages favor recombination of all taxa of Torrendia in Amanita.
The type species of Torrendia is T. pulchella Bres. (1902). See
Malençon (1955), Bas (1975), Miller and Horak (1992), Tulloss (2005b). The image of T. pulchella is a line drawing by
C. Bas that is used with his permission.
[NB: Images and well-documented
dried collections are sought by both
editors.] |
I. Subgenus Amanita
Taxa of this subgenus have
spores that do not darken when exposed to iodine
(e.g., when mounted in Melzer's Reagent) --
inamyloid spores. Section
names follow usage of
Corner and Bas (1962) and Bas (1969) as emended in Yang (1997).
      Section Amanita - Taxa of this section
have a primordium in which the developing cap and stipe take up
a part that is offset toward the top of the primordium and
disproportionately smaller than in the mature basidiome.
This usually results in the taxa having a prominently bulbous
base to their stipes at least prior to maturity or
aging. The type species of Amanita (Amanita muscaria) belongs to this
section. Almost all species in this section lack a membranous, saccate universal
veil (volva) enclosing the stipe's bulbous base. The few
taxa presently known to have such a universal veil are limited
to South America; however, similar taxa may be found (for example) in Africa,
Australia, and India. A number of the taxa in this section
contain compounds causing the Pantherine Syndrome in humans and
other animals.
       Section Vaginatae - Taxa of this section
have a primordium in which the stipe is centered vertically and
expands over its total length leaving no unexpanded bulb at the
stem base in mature material. A cupulate volva on the
stipe base should not be mistaken
for a bulb. Longitudinal sectioning of basidiomes can help
clarify the situation. While it is not a fundamental
defining character of this section, many of the taxa have
membranous saccate universal veils (volvas). The taxa that
lack such a universal veil are
distributed nearly world wide and have a universal veil that is
friable or submembranous.
The type species of this section is Amanita vaginata (Bull. : Fr.) Lam.
(1783 ["1784"]).
      Section Caesareae - Taxa assigned to
this section were considered to belong to section Vaginatae by Corner and Bas
(1962). The editors and other contemporary taxonomists
consider the group of taxa fitting the definition of section Vaginatae, but having a membranous
partial veil, to be assignable to the present taxon. It is
proposed that the type species of both Amarrendia and Torrendia both are
assignable to section Caesareae. A world key to the
species of section Caesareae is
available (over 540 Kb PDF) on this site (here).
The type species of this section is Amanita caesarea (Scop. : Fr.)
Pers. (1801).
II. Subgenus Lepidella
Taxa of this subgenus have
spores that darken (faintly to intensely) when exposed to iodine
(e.g., when mounted in Melzer's Reagent) -- amyloid spores. Section names
are used in the sense of Corner and Bas (1962) and Bas (1969) with
the exception of moving taxa of what is now sometimes called
subsect. Mappae (A. asteropus, A. brunnescens, A. bulbosa, A. porphyria, A. sinocitrina,
etc.) from their previous placement in sect. Phalloideae to the currently accepted
placement in sect. Validae.
      Section
Lepidella
- Taxa of this section include
some with the characters judged "most primitive" or "least
derived" in Amanita.
Unpublished molecular studies support the hypothesis these same
species are basal to the Amanita
evolutionary tree. Hence, from two very different
perspectives, there is some support for the idea that the
earliest Amanita we might today
consider to belong in the genus (were we to see it) would be an
entity we would also consider to belong in section Lepidella. The margin of the
pileus of species in this section is always appendiculate at
first. The stipe base very infrequently has a small and
rather weakly membranous limb arising from a basal bulb;
however, for the majority of taxa in the section, this is not
the case. Beyond the definition provided by the last two
sentences, there are a number of other characters that, while
not shared by all members of the section are notable field
characters that are often present. Outside of Australia
and Africa, few of these taxa have basidiome pigmentations other
than in the gray, brown, or black ranges. In fact, a
majority of the taxa are white or pallid. Taxa often have
flocculent material on them that comes off on the fingers of
collectors; the partial veil (annulus or ring) is often weakly
structured and may not persist to maturity of the basidiome; and
many taxa also exhibit an odor. These odors range widely
("anise" or "garlic" to "stale tiger urine"), but many are
rather unpleasant. While some species in this section may be edible, a growing number have
been found to contain amino acids that are severely toxic to the
human kidney and liver.
The type species of this section is Amanita vittadinii (Morreti) Vitt.
(1826).
    Section
Amidella - Taxa of this section
have an appendiculate cap margin combined with a robust,
multi-layered universal veil that is found in a mature material
as a saccate volva on a totally elongating stipe. The
innermost layer of the volva is often left on the pileus as fine
flocculence or as a thin layer looking like cracked paint.
Some Amanita "rules of thumb" such
as "if the pileus (cap) margin is striate and the short gills
(lamellulae) are squarely cut off (truncate), then the spores
will be inamyloid" are thoroughly violated by species of this
section. Morphological and molecular evidence both suggest
that this section arose from an ancestry among the
limbate-volva'd taxa of section Lepidella. The number of taxa
assignable to this section is rather small. They are
clustered into two or more groups. The type species is A. volvata. It is a North
American taxon with very similar taxa present in western Europe
and eastern Asia. This group of taxa often exhibit pink to
brick-colored bruising/staining reactions. A smaller
number of taxa (limited to Eurasia as far as is known) are pure
white except for yellow to tan developing on the exterior of the
volva. South America and a number of island regions (e.g.,
New Zealand) apparently lack taxa in this section. The
situation in Australia is not well understood, although several
taxa are probably assignable to section Amidella. It is possible that
some taxa there are modern representatives of taxa that were
intermediary between sections Lepidella and Amidella. Some older keys say
that gills of members of this section dry a very dark
color. Experimentation indicates that this is not true if
a specimen is dried quickly with relatively low ambient
humidity.
The
type species of this section is A. volvata Peck (Lloyd)
(1898).
     Section
Phalloideae - This section is defined
in part by the cap margin's lacking friable, appendiculate
material. All known species are annulate and all but one
[A. longitibiale Tulloss et al. (1995)] bear a limbate volva
attached to the top of the stipe's basal bulb. All known
species lack clamps at the bases of basidia. Pileus
coloration ranges widely from pure white to bright colors to the
pointilliste or marbled appearance seen in the type species --
A. phalloides (Fr. : Fr.) Link
(1833). Many, but not all, of the taxa in this section
contain one or more amatoxins and/or phalloidin. The
amatoxins are not destroyed by heat and are the principal cause
of death by Amanita poisoning.
Specimens thought to belong to this section, should be checked
for (1) a complexly layered volva that is saccate (rather than
limbate), which would suggest placement in section Amidella, or (2) a very thin, weak, and
(sometimes) short volval limb that may have a powdery inner
layer left on the mushroom's cap, which would suggest placement
in (for example) subsection Limbatulae Bas (1969) of section Lepidella.
     Section
Validae - This section is defined
in part by the cap margin's lacking friable, appendiculate
material and the absence of a membranous universal veil.
All known species are annulate. All known species lack
clamps at the bases of basidia. Pileus coloration ranges
widely from pure white to a variety of bright colors. Two
distinctive groups within the section exhibit striking staining
of the flesh when they are bruised or sectioned. Many of
the taxons of this section that have been tested contain a
hemolytic compound that is destroyed by heat. If material
is ingested uncooked, gastrointestinal distress with a rather
rapid onset may be experienced Hence, in several parts of
the world, species in this section are commonly eaten, but never ingested raw.
The type species
for this section is Agaricus validus
Fr. (1838) {now considered (by some authors) a posterior synonym
of Agaricus excelsus Fr. (1821) [= A.
excelsa (Fr. : Fr.) Bertill. (1866)]}.
Biogeography of Amanita
Publications (PDFs) and
links
Methodology for
Mophological Studies
- Glossary - from
Agriculture Canada site
- Tulloss' Form for Notes on Fresh Material [
]
- Recording Forms for
Macrochemical Spot Testing (t.b.d.)
- Form for
Phenoloxidase Spot Testing (developed by C. Marr, RET,
& A. Montoya-Esquivel) Blank form
(t.b.d.)
.
- Form for Phenoloxidase
Spot Testing filled in for a species of sect. Vaginatae (t.b.d.)
- Meaning of Biometric Variables
- a few terms used in Tulloss' descriptions of spores
and microscopic anatomy
Nomenclatorial studies (t.b.d)
-
Partial
bibliography -
Mostly Twentieth Century (worldwide) and originally based
on the bibliography in A Seminar on
Amanita (Tulloss, 1998). This is a work in
progress.
Extended
bibliography (t.b.d.)
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