Common puffball Lycoperdon perlatum, copyright Patrick Schifferli.

Belongs within: Agaricaceae.

When the wolf breaks wind
Published 21 December 2017
Common puffballs Lycoperdon perlatum, copyright H. Krisp.

In an earlier post, I described the way in which the ‘gasteromycetes’ of historical fungal classifications have come to be expunged as a category. The enclosure of spore-producing structures within a contained fruiting body such as a puffball, instead of exposed on a membrane such as on the underside of a mushroom cap, is something that has evolved many times in fungal history. One possible suggestion for why this may occur is as a protection against moisture loss, allowing the fungus to thrive in drier or more exposed habitats than before.

In that earlier post, I also mentioned off-hand that one of the best known groups of ‘gasteromycetes’, the puffballs of the Lycoperdaceae, are in fact close relatives of some of the best known typical mushrooms in the Agaricaceae. Indeed, it appears that recent authors may go so far as to synonymise the two families. Puffballs emerge as globular fruiting bodies that become packed with spores as they mature, until one or more openings develop in the external skin of the fruiting body and allow the pores to escape. Supposedly many puffballs are quite edible if collected before the spores begin to develop, though I’ve never tried myself. Particularly sought in this regard is the giant puffball Calvatia gigantea whose fruiting bodies grow particularly large; supposedly, examples have been found over a metre in diameter and weighing up to twenty kilogrammes.

Giant puffball Calvatia gigantea, copyright Alan Wolf.

Dispersal of spores from puffballs may be achieved in a number of ways. In species found in habitats with more regular rainfall, such as species of the genus Lycoperdon, spores are spread by ‘boleohydrochory’ (Gube & Dörfelt 2011). ‘-Chory’ means dispersal, ‘-hydro-‘ obviously means water, ‘boleo-‘ I think may mean something like ‘throw’. The puffball opens through a hole in the top, and drops of rain (or other sources of pressure such as being tapped by an animal) cause a puff of spores to be squeezed out. The water may then carry the spores away. The name Lycoperdon, as it happens, literally translates as ‘wolf fart’, and this is another one of those names I am completely at a loss to explain. The ‘fart’, obviously, refers to the appearance of the spore puffs, but what on earth do they have to do with wolves?

Tumbling puffballs Bovista pila, copyright Dan Molter.

Other puffballs may spread their spores via ‘anemochory’, dispersal by wind. This is particularly the case with species found in drier habitats. Some species, such as some members of the genus Bovista, exhibit a variation on this called ‘geanemochory’ in which the entire puffball becomes detached and blown about by the wind, with the spores escaping through openings in the external shell like pepper being shaken from a pepper-pot. Differences in dispersal method between puffball species are generally reflected by differences in their spore morphology. Hydrochorous species usually have strong ornamentation, with the outside of the spore being covered with warts or the like. These warts provide more surface area for the water to catch onto; they may also help prevent the spores from clumping together. In contrast, anemochorous species have spores that are smooth, making them more streamlined for being blown through the air or, particularly in the case of geanemochorous species, making them less likely to become trapped by hyphae or other structures inside the fruiting body itself and so facilitating their escape.

Systematics of Lycoperdales
Lycoperdales [Lycoperdeae]
| i. s.: Geasterites Pia 1927T93, KC01
| `--G. florissantensis Cockerell 1908T93
|--Mycenastrum Desvaux 1842C27 (see below below for synonymy)
| `--M. corium (Guersent) Desvaux 1842 (see below for synonymy)C27
| i. s.: Abstoma Cunningham 1926KC01
| `--*A. purpureum (Lloyd) Cunningham 1927 [=Catastoma purpureum Lloyd 1922]C27
| Glyptoderma Heim & Perr.-Bertr. 1971 [incl. Acutocapillitium Ponce de León 1976]KC01
| Arachnion Schwein. 1822 [incl. Scoleciocarpus Berk. 1843]KC01
| `--A. lloydianumKB01
| ‘Arachniopsis’ Long 1917 non Spruce 1882KC01
| Calbovista Morse ex Seidl 1995KC01
| Calvatiopsis Hollós 1929KC01
| Disciseda Czernaiaiev 1845KC01 [incl. Bovistina Long & Stouffer 1941KC01, Catastoma Morgan 1892C27]
| |--D. anomalaC27
| |--D. candida (Schweinitz) Lloyd 1902 (see below for synonymy)C27
| |--D. hyalotrixC27
| |--D. hypogaeaC27
| |--‘Catastoma’ magnum Lloyd 1917 (n. d.)C27
| |--D. subterranea [=Catastroma subterraneum]KC01
| `--D. verrucosa Cunningham 1927C27
| Lycogalopsis Fisch. 1886 [incl. Enteromyxa Ces. 1879]KC01
| Japonogaster Kobayasi 1989KC01
| Lycoperdopsis Henn. 1899KC01
|--Calvatia Fries 1849 (nom. cons.)KB01, C27 (see below for synonymy)
| |--C. caelata (Bulliard) Morgan 1890 (see below for synonymy)C27
| |--C. complutensisKB01
| |--C. fragilisKB01
| |--C. gigantea (Persoon) Cunningham 1927C27 (see below for synonymy)
| `--C. lilacina (Berkeley & Montagne) Lloyd 1905 (see below for synonymy)C27
`--+--‘Lycoperdon’ pyriforme Schaeffer ex Persoon 1801KB01, C27
`--+--Bovista Dillenius ex Persoon 1801KB01, C27 (see below for synonymy)
| | i. s.: B. bovistoidesSPB07
| | B. brunnea Berkeley 1855C27
| | B. cunninghamii Kreisel 1967SPB07
| | B. fulvaSPB07
| | B. limosaKB01
| | B. nigrescensKB01
| | B. oblongispora (Lloyd) Bottomley 1948 [=Bovistella oblongispora Lloyd 1917]SPB07
| | B. ovalispora Cooke & Massee 1887 (n. d.)C27
| | B. purpurea Lloyd 1923C27
| | B. trachyspora [=Lycoperdon trachyspora]SPB07
| |--+--B. paludosaKB01
| | `--B. plumbeaKB01
| `--+--B. polymorphaKB01
| `--B. pusilla Persoon 1801KB01, C27 (see below for synonymy)
`--Lycoperdon Tournefort ex Persoon 1801KB01, C27 (see below for synonymy)
| i. s.: L. compactum Cunningham 1927C27
| L. cruciatumKC01
| L. depressum Bonorden 1857C27
| L. gemmatumC96
| L. glabrescens Berkeley 1860 (see below for synonymy)C27
| L. hiemaleC27
| L. hoyleiC27
| L. mixtecorumKC01
| L. oblongisporumSPB07
| L. polymorphum Vittadini 1842 (see below for synonymy)C27
| L. reticulatum (n. d.)C27
| L. scabrum (Lloyd) Cunningham 1927 (see below for synonymy)C27
| L. spadiceum Persoon 1809 [incl. L. cookei Massee 1887]C27
| L. sub-pratenseC27
|--+--Morganella Zeller 1948KB01, KC01
| | `--M. subincarnataKB01
| |--Vascellum Smarda 1958KB01, KC01
| | `--V. pratenseKB01 [=Lycoperdon pratenseC27]
| `--+--*L. perlatum Persoon 1801KB01, C27 (see below for synonymy)
| | |--L. p. var. perlatumKB01
| | `--L. p. var. bonordeniiKB01
| `--L. marginatumKB01
`--+--Bovistella Morgan 1892KB01, C27 [incl. Calvatiella Chow 1936KC01]
| |--B. bovistoides (Cooke & Massee) Lloyd 1906 (see below for synonymy)C27
| |--B. cuprica Lloyd 1915 (n. d.)C27
| `--B. radicataKB01
`--+--+--L. echinatumKB01
| `--L. mammiformeKB01
`--+--+--L. foetidumKB01
| `--L. umbrinumKB01
`--+--L. lividumKB01
`--L. molleKB01

Bovista Dillenius ex Persoon 1801KB01, C27 [incl. Piemycus Raf. 1813KC01, Piesmycus Raf. 1808KC01, Pseudolycoperdon Velen. 1947KC01, Sackea Rostk. 1844KC01]

Bovista pusilla Persoon 1801KB01, C27 [=Lycoperdon pusillum Persoon 1809C27; incl. L. dermoxanthum Vitt. 1842C27, Bovista dermoxanthum (Vitt.) de Toni in Sacc. 1888C27, L. microspermum Berkeley in Hooker 1854C27, L. mundulum Kalchbr. 1880C27, Bovista mundula (Kalchbr.) de Toni in Sacc. 1888C27, L. pseudopusillum Hollos 1903C27, L. semi-immersum Lloyd 1924C27]

Bovistella bovistoides (Cooke & Massee) Lloyd 1906 [=Mycenastrum bovistoides Cooke & Massee 1888, Scleroderma bovistoides (Cooke & Massee) de Toni in Sacc. 1888]C27

Calvatia Fries 1849 (nom. cons.)KB01, C27 [incl. Bovistaria (Fr.) Karst. 1889KC01, Eriosphaera De Toni 1888 non Dietr. 1817KC01, Gastropila Homrich & Wright 1973KB01, KC01, Handkea Kreisel 1989KC01, Hippoperdon Mont. 1842 (nom. rej.)KC01, Hypoblema Lloyd 1902KC01, Langermannia Rostk. 1839KB01, KC01, Lanopila Fr. 1848KC01, Lasiosphaera Reichardt 1870KC01, Omalycus Raf. 1814KC01, Pila Speg. 1923 nec Klein 1758 (ICZN) nec Röding 1798 (ICZN) nec Bertrand & Renault 1892KC01]

Calvatia caelata (Bulliard) Morgan 1890 [=Lycoperdon caelatum Bulliard 1809; incl. L. favosum Bon. 1857, Calvatia favosa (Bon.) Lloyd 1905, L. fontanesii Dur. & Montagne 1849, C. fontanesii (Dur. & Montagne) Lloyd 1905, L. sinclairii Berkeley 1887, C. sinclairii (Berkeley) Lloyd 1905]C27

Calvatia gigantea (Persoon) Cunningham 1927C27 [=Lycoperdon giganteum Persoon 1801C27, Bovista gigantea (Persoon) Nees 1817C27, Langermannia giganteaKC01; incl. Lycoperdon bovista Fries 1829C27, Calvatia maxima (Schaeff.) Morg. 1890C27, C. primitiva Lloyd 1905C27]

Calvatia lilacina (Berkeley & Montagne) Lloyd 1905 [=Bovista lilacina Berkeley & Montagne in Hooker 1845, Lycoperdon lilacinum (Berkeley & Montagne) Massee 1887; incl. Calvatia cyathiformis (Bosc.) Morgan 1890, L. novae-zelandiae Lev. 1846]C27

Disciseda candida (Schweinitz) Lloyd 1902 [=Bovista candida Schweinitz 1822; incl. B. circumscissa Berkeley & Curtis 1874, Catastoma circumscissa (Berkeley & Curtis) Morgan 1892, Disciseda circumscissa (Berkeley & Curtis) Hollos 1902]C27

Lycoperdon Tournefort ex Persoon 1801KB01, C27 [=Utraria Quél. 1873KC01; incl. Capillaria Velen. 1947 nec Roussel 1806 nec Pers. 1822 KC01, Cerophora Raf. 1808 KC01, Globaria Quél. 1876C27, Priapus Raf. 1808 non Linnaeus 1758 (ICZN)KC01, Sufa Adans. 1763KC01, Ultraria Quel. 1876C27]

Lycoperdon glabrescens Berkeley 1860 [=Bovistella glabrescens (Berkeley) Lloyd 1905; incl. B. australiana Lloyd 1905, B. rosea Lloyd 1906]C27

*Lycoperdon perlatum Persoon 1801KB01, C27 [incl. L. colensoi Cooke & Massee 1887C27, L. excipuliforme (Scopoli) Vitt. 1842C27, L. macrogemmatum Lloyd 1906C27, L. montanum Quel. 1876C27, L. tasmanicum Massee 1901C27]

Lycoperdon polymorphum Vittadini 1842 [incl. L. cepaeforme (Bull.) Massee 1887, L. furfuraceum Schaeff. ex de Toni 1888, L. hungaricum Hollos 1901, L. nigrum Lloyd 1905]C27

Lycoperdon scabrum (Lloyd) Cunningham 1927 [=Bovistella scabra Lloyd 1906; incl. B. nigrica Lloyd 1922]C27

Mycenastrum Desvaux 1842C27 [incl. Endonevrum Czern. 1845KC01, Pachyderma Schulzer 1876 non Blume 1826KC01; MycenastraceaeKB01]

Mycenastrum corium (Guersent) Desvaux 1842 [=Lycoperdon corium Guersent in de Candolle 1815, Scleroderma corium Grav. in Duby 1830; incl. Mycenastrum olivaceum Cooke & Massee 1887, Scleroderma olivaceum (Cooke & Massee) de Toni in Sacc. 1888, M. phaeotrichum Berkeley in Hooker 1843, S. phaeotrichum (Berkeley) de Toni in Sacc. 1888, M. spinulosum Peck 1883]C27

*Type species of generic name indicated


[C96] Campbell, N. A. 1996. Biology 4th ed. The Benjamin/Cummings Publishing Company, Inc.: Menlo Park (California).

[C27] Cunningham, G. H. 1927. Lycoperdaceae of New Zealand. Transactions and Proceedings of the New Zealand Institute 57: 187–217.

Gube, M., & H. Dörfelt. 2011. Gasteromycetation in Agaricaceae s. l. (Basidiomycota): morphological and ecological implementations. Feddes Repertorium 122 (5–6): 367–390.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).

[KB01] Krüger, D., M. Binder, M. Fischer & H. Kreisel. 2001. The Lycoperdales. a molecular approach to the systematics of some gasteroid mushrooms. Mycologia 93 (5): 947–957.

[SPB07] Sharma, J. R., K. N. Pandey & D. Bisht. 2007. Two new records of the genus Bovista Pers.: Pers. (Gasteromycetes) from India. Bulletin of the Botanical Survey of India 49: 225–230.

[T93] Taylor, T. N. 1993. Fungi. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 9–13. Chapman & Hall: London.

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