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Studies in the Genus Amanita Pers.
(Agaricales, Fungi)
                                        ( contact us )





These pages are dedicated to

Dr. Cornelis Bas
Leiden, The Netherlands

He transformed the study of Amanita.


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Site Navigation

- Bibliography (Partial)

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Editors

Dr. Rodham E. Tulloss*
Herbarium Rooseveltensis Amanitarum
P. O. Box 57, Roosevelt, New Jersey 08555-0057, USA
[ *Research Assoc. (hons.), New York Botanical Garden ]

Prof. Dr. Zhu-liang Yang
Key Laboratory of Biodiversity and Biogeography
Kunming Institute of Botany, Chinese Academy of Sciences
Kunming, Yunnan 650204, China


The New York Botanical GardenKunming Institute of Botany, Yunnan Prov., ChinaPersooniaINBio, Costa Rica Great Smoky Mountains ATBI EOL

OUR THANKS: We greatly appreciate permission granted to include on the pages of this site line illustrations of C. Bas and water colors of J. E. B Corner that originally appeared in the journal Persoonia.  We are very grateful for the honor of having had material from this site used in the original sample pages (May, 2007) of the Encyclopedia of Life.  Tulloss is very grateful for the opportunities of working with the Great Smoky Mountains ATBI and with the Fungus Taxonomic Working Group del Instituto Nacional de Biodiversidad de Costa Rica.

NOTICE: This site is not associated in anyway with sites selling wild mushrooms in any form, the extracts of such mushrooms, or any related paraphernalia. Such sites have made unauthorized use of our pages to create an apparent relation to scientific research; however, the visitor to those sites should be suspicious of out-of-date pages with altered design or altered background colors. Pages stolen from this site and bearing an editor's name or even a contributor's copyright notice are not authorized by the Amanita Studies site's editors in anyway. We do not promote or encourage experimentation with ingestion of toxic Amanita species.

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Bibliography (Partial)

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Amanita crocea Amanita crocea


Bibliography (Partial)

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Molecular Phylogeny

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Search this Site

Site News: 
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Amanita crocea Amanita crocea


Bibliography (Partial)

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Molecular Phylogeny

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Search this Site

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Amanita crocea Amanita crocea


Bibliography (Partial)

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Search this Site

Site News: 
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Amanita crocea Amanita crocea


Bibliography (Partial)

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Search this Site

Site News: 
Check for updates!


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Amanita crocea Amanita crocea


Bibliography (Partial)

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Nomenclatorial Studies

Search this Site

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Check for updates!


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Amanita crocea Amanita crocea


Bibliography (Partial)

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Molecular Phylogeny

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Search this Site

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Amanita crocea Amanita crocea


Bibliography (Partial)

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Molecular Phylogeny

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Nomenclatorial Studies

Search this Site

Site News: 
Check for updates!


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Amanita crocea Amanita crocea


Biogeography of Amanita


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Molecular Phylogeny

Mushroom Poisoning (general)


Nomenclatorial Studies

Search this Site

Site News: 
Check for updates!


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Amanita crocea Amanita crocea


Bibliography (Partial)

Biogeography of Amanita


Keys & Checklists/
Picturebooks


Methodology  (Morphological Studies)

Molecular Phylogeny

Mushroom Poisoning (general)


Nomenclatorial Studies

Search this Site

Site News:
Check for updates!


Species Pages

Toxins of Amanita


Amanita crocea Amanita crocea



Introduction: Welcome to the Amanita Studies site.  This page and subordinate pages provide data on species of the macrofungal, largely ectomycorrhizal genus Amanita and the family Amanitaceae (Agaricales, Basidiomycetes, Fungi) to which it belongs.  Text is 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 in Amanita sect. Vaginatae).

At present, the section level lists include 566 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" (although some descriptions need further work) for 518 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 [further] developed"]

Background tile depicts A. muscaria subsp. flavivolvata of North and Central America.
Tiles in navigation column (at left) depict A. crocea of Europe.


NEW

Search Amanita studies site:

AND and OR functions work.
All capabilities and syntax of queries are those of Google.

This free script provided by JavaScript Kit

Having trouble finding something?

The greatest search problem of our users is misspelling of names.
Try using the sectional lists.

Another source of informative links is the
dmoz Open Directory Project's Amanitaceae category

dmoz Open Directory Project's Amanita category


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.  NEW

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 of Amarrendia are sought by both editors.]


Limacella illinita - Tlaxcala, Mexico - RET Limacella glischra - Chiricahua Mtns., Arizona, USA - RET Limacella illinita - Tlaxcala, Mexico - RET Limacella glischra - Chiricahua Mtns., Arizona, USA - RET Limacella illinita - Tlaxcala, Mexico - RET Limacella glischra - Chiricahua Mtns., Arizona, USA - RET Limacella illinita - Tlaxcala, Mexico - RET 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 pulchella - C. BasTorrendia pulchella - C. BasTorrendia pulchella - C. BasTorrendia pulchella - C. BasTorrendia pulchella - C. BasTorrendia pulchella - C. BasTorrendia pulchella - C. BasTorrendia pulchella - C. BasTorrendia pulchella - C. BasTorrendia pulchella - C. BasTorrendia pulchella - C. BasTorrendia - 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 of Torrendia 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 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. : 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 [ FOR YOUR USE ]
  • 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.)

Nomenclatorial studies (t.b.d)

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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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Last updated 10 October 2009.
This page is maintained by R. E. Tulloss.
Copyright 1999, 2000, 2001, 2002, 2003, 2004, 2005, 2006, 2007, 2008, 2009 by Rodham E. Tulloss.
Photographs copyright 2002 by Rodham E. Tulloss.

Development support for these pages was provided by Lindsay Possiel during the summers of 2004, 2005, and 2006.
The editors express their sincere gratitude for a job very well done.