Editing Puffball

{{Short description|Fungus}}
{{Other uses}}
{{Multiple image
 | align = right
 | direction = vertical
 | width = 200
 | image1 = Flaschenstäubling.jpg
 | caption1 = ''[[Lycoperdon perlatum]]''
 | image2 = Calbovista subsculpta 45731.jpg
 | caption2  = ''[[Calbovista subsculpta]]''
 | image3 = Calvatia cyathiformis 3.JPG
 | caption3 = ''[[Calvatia cyathiformis]]''
 | footer = Puffballs are found in several genera of the division [[Basidiomycota]].
}}

'''Puffballs''' are a type of [[fungus]] featuring a ball-shaped [[fruit body]] that (when mature) bursts on contact or impact, releasing a cloud of dust-like spores into the surrounding area. Puffballs belong to the division [[Basidiomycota]] and encompass several genera, including ''[[Calvatia]]'', ''[[Calbovista]]'' and ''[[Lycoperdon]]''.<ref>{{Cite web|url=http://www.mssf.org/cookbook/puffballs.html|title=Wild About Mushrooms:Puffballs|last1=Freedman|first1=Louise|last2=Mycological Society of San Francisco|author2-link=Mycological Society of San Francisco|year=2000|orig-year=1987|archive-url=https://web.archive.org/web/20080928081858/http://www.mssf.org/cookbook/puffballs.html|archive-date=2008-09-28|url-status=live|access-date=2008-08-03}}</ref> The puffballs were previously treated as a taxonomic group called the [[Gasteromycetes]] or Gasteromycetidae, but they are now known to be a [[polyphyletic]] assemblage.

The distinguishing feature of all puffballs is that they do not have an open cap with spore-bearing [[lamella (mycology)|gills]]. Instead, [[spore]]s are produced internally, in a spheroidal fruit body called a ''gasterothecium'' (gasteroid 'stomach-like' [[basidiocarp]]). As the spores mature, they form a mass called a [[gleba]] in the centre of the fruitbody that is often of a distinctive color and texture. The basidiocarp remains closed until after the spores have been released from the [[basidia]]. Eventually, it develops an aperture, or dries, becomes brittle, and splits, and the spores escape. The spores of puffballs are ''statismospores'' rather than ballistospores, meaning they are not forcibly extruded from the basidium. Puffballs and similar forms are thought to have [[convergent evolution|evolved convergently]] (that is, in numerous independent events) from [[Hymenomycetes]] by ''gasteromycetation'', through [[secotioid]] stages. Thus, 'Gasteromycetes' and 'Gasteromycetidae' are now considered to be descriptive, morphological terms (more properly ''gasteroid'' or ''gasteromycetes'', to avoid taxonomic implications) but not valid [[cladistic]] terms.

True puffballs do not have a visible stalk or stem, while stalked puffballs do have a stalk that supports the gleba. None of the stalked puffballs are edible as they are tough and woody mushrooms.<ref name="MushroomsNA"/> The [[Hymenogastrales]] and ''[[Enteridium lycoperdon]]'', a [[slime mold]], are the ''false puffballs''. A gleba which is powdery on maturity is a feature of true puffballs, stalked puffballs and earthstars. False puffballs are hard like rock or brittle. All false puffballs are inedible, as they are tough and bitter to taste. The genus ''[[Scleroderma (fungus)|Scleroderma]]'', which has a young purple gleba, should also be avoided.<ref name="MushroomsNA">{{Cite book
  | last = Miller
  | first = Orson K.
  | title = Mushrooms of North America
  | place = New York, NY
  | publisher = E.P. Dutton Publisher
  | year = 1977
  | page = 298
  | isbn = 978-0-525-47482-1}}</ref>

Puffballs were traditionally used in [[Tibet]] for making [[ink]] by burning them, grinding the ash, then putting them in water and adding glue liquid and "a {{transliteration|bo|nye shing ma}} decoction", which, when pressed for a long time, made a black dark substance that was used as ink.<ref>Cuppers, Christoph (1989). "On the Manufacture of Ink." ''Ancient Nepal – Journal of the Department of Archaeology'', Number 113, August–September 1989, p. 5.</ref> Rural Americans burned the common puffball with some kind of [[bee smoker]] to anesthetize [[honey bee]]s as a means to safely procure honey; the practice later inspired experimental medicinal application of the puffball smoke as a surgical general anesthetic in 1853.<ref>{{Cite journal|date=2021-06-01|title=A brief history of carbon monoxide and its therapeutic origins|url=https://www.sciencedirect.com/science/article/abs/pii/S1089860321000367|journal=Nitric Oxide|language=en|volume=111-112|pages=45–63|doi=10.1016/j.niox.2021.04.001|issn=1089-8603|last1=Hopper|first1=Christopher P.|last2=Zambrana|first2=Paige N.|last3=Goebel|first3=Ulrich|last4=Wollborn|first4=Jakob|pmid=33838343|s2cid=233205099|access-date=2021-04-26|archive-date=2021-04-27|archive-url=https://web.archive.org/web/20210427035112/https://www.sciencedirect.com/science/article/abs/pii/S1089860321000367|url-status=live}}</ref>

==Edibility and identification==
[[File:Puffball fungai.jpg|thumb|upright=1|Cloud of spores poofs out of a puffball fungus]]

While most puffballs are not poisonous, some often look similar to young [[Agaricales|agarics]], and especially the deadly [[Amanita]]s, such as the [[death cap]] or [[destroying angel]] mushrooms. Young puffballs in the edible stage, before maturation of the gleba, have undifferentiated white flesh within, whereas the gills of immature Amanita mushrooms can be seen if they are closely examined. They can be very toxic.

[[File:Puffball Mushrooms On Sale.jpg|thumb|upright=1|Puffball mushrooms on sale at a market in England]]

The [[giant puffball]], ''Calvatia gigantea'' (earlier classified as ''Lycoperdon giganteum''), reaches {{convert|1|ft}} or more in diameter, and is difficult to mistake for any other fungus. It has been estimated that, when mature, a large specimen of this fungus will produce around [[Orders of magnitude (numbers)#1012|7 × 10{{sup|12}}]] spores, which is more than any other known organism.

Not all true puffball mushrooms are without stalks.{{Inconsistent|date=November 2021|reason=Lead section states true puffballs have no stems}} Some may also be stalked, such as the ''[[Podaxis pistillaris]]'', which is also called the "false shaggy mane". There are also a number of false puffballs that look similar to the true ones.<ref name="MushroomsNA"/>

[[File:Common puffball, releasing spores in a burst by compressing the body.webm|thumb|Common puffball, releasing spores in a burst by compressing the body]]

===Stalked===
Stalked puffballs species:<ref name="MushroomsNA"/>
* ''[[Battarrea phalloides]]''
* ''[[Calostoma cinnabarina]]'' (Stalked Puffball-in-Aspic)<ref name="expert">{{cite web
 |last         = Kuo
 |first        = M.
 |title        = Puffballs
 |publisher    = MushroomExpert.Com
 |date         = February 2006
 |url          = http://www.mushroomexpert.com/puffballs.html
 |access-date   = 2008-08-03
 |archive-url  = https://web.archive.org/web/20080921103804/http://www.mushroomexpert.com/puffballs.html
 |archive-date = 2008-09-21
 |url-status     = live
}}</ref>
* ''[[Pisolithus tinctorius]]''
* ''[[Tulostoma]]'' (genus)

===True===
True puffballs genera and species:<ref name="MushroomsNA"/>
* ''[[Bovista]]'' – various species, including:
** ''[[Bovista aestivalis]]''
** ''[[Bovista dermoxantha]]''
** ''[[Bovista nigrescens]]''
** ''[[Bovista plumbea]]''
* ''[[Calvatia]]'' – various species, including:
** ''[[Calvatia bovista]]''
** ''[[Calvatia craniiformis]]''
** ''[[Calvatia cyathiformis]]''<ref name="MushroomsNA"/><ref name="expert"/>
** ''[[Calvatia gigantea]]''<ref name="MushroomsNA"/><ref name="expert"/>
** ''[[Calvatia booniana]]''
** ''[[Calvatia fumosa]]''
** ''[[Calvatia lepidophora]]''
** ''[[Calvatia pachyderma]]''
** ''[[Calvatia sculpta]]''
** ''[[Calvatia subcretacea]]'' – edible<ref>{{Cite book|last1=Davis|first1=R. Michael|url=https://www.worldcat.org/oclc/797915861|title=Field Guide to Mushrooms of Western North America|last2=Sommer|first2=Robert|last3=Menge|first3=John A.|publisher=[[University of California Press]]|year=2012|isbn=978-0-520-95360-4|location=Berkeley|pages=366–367|oclc=797915861|access-date=2021-06-29|archive-date=2022-06-03|archive-url=https://web.archive.org/web/20220603052838/https://www.worldcat.org/title/field-guide-to-mushrooms-of-western-north-america/oclc/797915861|url-status=live}}</ref>
** ''[[Calbovista subsculpta]]''
* ''[[Handkea]]'' – various species, including:
** ''[[Handkea utriformis]]''
* ''[[Lycoperdon]]'' – various species, including:
** ''[[Lycoperdon candidum]]''
** ''[[Lycoperdon echinatum]]''
** ''[[Lycoperdon fusillum]]''
** ''[[Lycoperdon umbrinum]]''
* ''[[Scleroderma (fungus)|Scleroderma]]'' – various species, including:
** ''[[Scleroderma auratium]]''
** ''[[Scleroderma geaster]]'' – not edible<ref>{{cite book |last=Phillips |first=Roger |title=Mushrooms and Other Fungi of North America |url=https://archive.org/details/mushroomsotherfu0000phil |url-access=registration |year=2010 |publisher=Firefly Books |location=Buffalo, NY |isbn=978-1-55407-651-2 |page=[https://archive.org/details/mushroomsotherfu0000phil/page/335 335]}}</ref>

===False===
False puffballs species:<ref name="MushroomsNA"/>
** ''[[Endoptychum agaricoides]]''
** ''[[Nivatogastrium nubigenum]]''
** ''[[Podaxis pistillaris]]''
** ''[[Rhizopogon rubescens]]''
** ''[[Truncocolumella citrina]]''

==Classification==
[[File:Lycoperdon pyriforme Sasata scale.jpg|thumb|right|''Apioperdon pyriforme'']]
[[File:Lycoperdon echinatum - Vesse de loup hérissée.JPG|thumb|right|''Lycoperdon echinatum'']]

Major orders:
* '''[[Agaricales]]''' (including now-obsolete orders [[Lycoperdales]], [[Tulostomatales]], and [[Nidulariales]])
** Basidiomycetes: Agaricales: Lycoperdaceae: ''Calvatia''
*** ''[[Calvatia booniana]]''<ref name="MushroomsNA"/>
*** ''[[Calvatia bovista]]'' (''[[Handkea utriformis]]'')
*** ''[[Calvatia craniiformis]]''<ref name="expert"/>
*** ''[[Calvatia cyathiformis]]''<ref name="MushroomsNA"/><ref name="expert"/>
*** ''[[Calvatia fumosa]]'' (Handkea fumosa)<ref name="MushroomsNA"/>
*** ''[[Calvatia gigantea]]''<ref name="MushroomsNA"/><ref name="expert"/>
*** ''[[Calvatia lepidophora]]''<ref name="MushroomsNA"/>
*** ''[[Calvatia rubroflava]]''<ref name="expert"/>
*** ''[[Calvatia sculpta]]''<ref name="MushroomsNA"/>
*** ''Calvatia subcretacea'' (''Handkea subcretacea'')<ref name="MushroomsNA"/>
** Basidiomycetes: Agaricales: Lycoperdaceae: ''Lycoperdon''
*** ''[[Lycoperdon foetidum]]'' (''[[Lycoperdon nigrescens]]'')
*** ''[[Lycoperdon perlatum]]''<ref name="expert"/>
*** ''[[Lycoperdon pulcherrimum]]''<ref name="expert"/>
*** ''[[Lycoperdon pusillum]]''
*** ''[[Lycoperdon pyriforme]]''
** Basidiomycetes: Agaricales: Lycoperdaceae: ''Vascellum''
*** ''[[Vascellum curtisii]]''<ref name="expert"/>
*** ''[[Vascellum pratense]]'' – edible when interior is white<ref>{{cite book |last=Phillips |first=Roger |title=Mushrooms and Other Fungi of North America |url=https://archive.org/details/mushroomsotherfu0000phil |url-access=registration |year=2010 |publisher=Firefly Books |location=Buffalo, NY |isbn=978-1-55407-651-2 |page=[https://archive.org/details/mushroomsotherfu0000phil/page/333 333]}}</ref>
* '''[[Geastrales]]''' and '''[[Phallales]]''' (related to '''[[Cantharellales]]'''),
** Basidiomycetes: Phallales: Geastraceae: ''Geastrum''
*** ''[[Geastrum coronatum]]''
*** ''[[Geastrum fornicatum]]''
*** ''[[Geastrum saccatum]]''<ref name="expert"/>
* '''[[Sclerodermatales]]''' (related to '''[[Boletales]]''')
** Basidiomycetes: Boletales: Sclerodermataceae: ''Scleroderma''
*** ''[[Scleroderma areolatum]]''<ref name="expert"/>
*** ''[[Scleroderma bovista]]''<ref name="expert"/>
*** ''[[Scleroderma cepa]]''
*** ''[[Scleroderma citrinum]]''<ref name="expert"/>
*** ''[[Scleroderma meridionale]]''<ref name="expert"/>
*** ''[[Scleroderma michiganense]]''<ref name="expert"/>
*** ''[[Scleroderma polyrhizum]]''<ref name="expert"/>
*** ''[[Scleroderma septentrionale]]''<ref name="expert"/>
* Various false-truffles ([[hypogaeic gasteromycetes]]) related to different hymenomycete orders

Similarly, the true truffles ([[Tuberales]]) are gasteroid [[Ascomycota]]. Their ascocarps are called tuberothecia.

==See also==
* [[Lycoperdonosis]]

==Footnotes==
{{Reflist|30em}}

==References==
* [http://tolweb.org/tree?group=Homobasidiomycetes&contgroup=Hymenomycetes Homobasidiomycetes] at the Tree of Life Web Project

==External links==
{{Commons category|Puffballs|position=left}}

{{Authority control}}

[[Category:Puffballs| ]]
[[Category:Edible fungi]]
[[Category:Fungus common names]]
[[Category:Basidiomycota]]
[[Category:Mushroom types]]