{{Short description|Species of puffball mushroom}} {{stack begin}} {{Speciesbox | image = Lycoperdon echinatum 54507 crop.jpg | image_alt = | image_caption = Mature specimen with brown spines | status = | status_system = | status_ref = | genus = Lycoperdon | species = echinatum | authority = Pers. (1797) | synonyms = *''Lycoperdon gemmatum'' var. ''echinatum'' <small>(Pers.) Fr. (1829)</small> *''Lycoperdon Hoylei'' <small>Berk. & Broome (1871)</small> *''Utraria echinata'' <small>(Pers.) Quél. (1873)</small> *''Lycoperdon americanum'' <small>Demoulin (1972)</small> | synonyms_ref = }} {{mycomorphbox | name = ''Lycoperdon echinatum'' | hymeniumType=gleba | stipeCharacter=NA | ecologicalType=saprotrophic | sporePrintColor=brown | howEdible=edible | howEdible2=inedible }} {{stack end}}
'''''Lycoperdon echinatum''''', commonly known as the '''spiny puffball''' or the '''spring puffball''', is a type of puffball mushroom in the family Agaricaceae. The saprobic species has been found in Africa, Europe, Central America, and North America, where it grows on soil in deciduous woods, glades, and pastures. It has been proposed that North American specimens be considered a separate species, ''Lycoperdon americanum'', but this suggestion has not been followed by most authors. Molecular analysis indicates that ''L. echinatum'' is closely related to the puffball genus ''Handkea''.
The fruit bodies of ''L. echinatum'' are {{convert|2|–|4|cm|in|1|abbr=on}} wide by {{convert|2|–|3.5|cm|in|1|abbr=on}} tall, supported by a small base, and densely covered with spines that are up to {{convert|0.6|cm|in|1|abbr=on}} long. The spines can fall off in maturity, leaving a net-like pattern of scars on the underlying surface. Initially white in color, the puffballs turn a dark brown as they mature, at the same time changing from nearly round to somewhat flattened. Young specimens of ''L. echinatum'' resemble another edible spiny puffball, ''Lycoperdon pulcherrimum'', but the latter species does not turn brown as it ages. The fruit bodies are edible when young, when the interior is white and firm and before it has turned into a powdery brown mass of spores. Laboratory tests have shown that extracts of the fruit bodies can inhibit the growth of several bacteria that are pathogenic to humans.
==Taxonomy and phylogeny== The species was first described by Christian Hendrik Persoon in 1797.<ref name=Fries1797/> It was later reduced to a variety of ''Lycoperdon gemmatum'' (as ''L. gemmatum'' var. ''echinatum''; ''L. gemmatum'' is now known as ''Lycoperdon perlatum''<ref name="urlIndex Fungorum - Synonymy for Lycoperdon gemmatum"/>) by Elias Magnus Fries,<ref name=Fries1829/> but American mycologist Charles Horton Peck, who extensively studied the North American distribution of the genus, raised it again to species level in 1879. He thought it worthy of status as a species distinct from ''L. gemmatum'' because of the different character of its warts, its much spinier appearance, and the smoother surface of the peridium underneath the spines.<ref name=Peck1879/> Miles Joseph Berkeley and Christopher Edmund Broome wrote of the fungus in 1871, but believed their specimen, collected from Reading, Berkshire, by Hoyle, represented a new species, which they called ''Lycoperdon Hoylei''. They wrote that their specimen agreed "exactly with an authentic specimen of Persoon's ''L. echinatum'' externally, who could, however, scarcely have overlooked the lilac spores."<ref name=Berkeley1871/> Despite the apparent difference in spore color, ''L. Hoylei'' is currently considered synonymous with ''L. echinatum''.<ref name="urlMycoBank: Lycoperdon hoylei"/> ''Utraria echinata'', named by Lucien Quélet in 1873,<ref name=Quelet1873/> is another synonym for ''L. echinatum''.<ref name="urlMycoBank: Utraria echinata"/>
In 1972, Vincent Demoulin described the species ''Lycoperdon americanum'' on the basis of a specimen found in North Carolina.<ref name=Demoulin1972/> Although he believed it to be a unique species,<ref name=Demoulin1973/> several authors consider it synonymous with ''L. echinatum''.<ref name=Abel1993/><ref name=Bessette2007/><ref name=Roody2003/><ref name=Kuo2003/> Phylogenetic analysis of the sequence and secondary structure of the ribosomal RNA (rRNA) genes coding for the internal transcribed spacer units suggests that ''Lycoperdon echinatum'' forms a clade with the puffball genus ''Handkea'', separate from the type species of ''Lycoperdon'', ''Lycoperdon perlatum''.<ref name="Krüger2008"/> In previous analyses that used only the rRNA sequences for phylogenetic comparison, ''L. echinatum'' formed a clade with ''L. mammiforme'', ''L. foetidum'', and ''Bovistella radicata'' (now known as ''Lycoperdon radicatum''<ref name="urlFungorum - Synonymy for Lycoperdon radicatum"/>), but separate from ''L. pyriforme''.<ref name="Krüger2001"/><ref name="Krüger2003"/>
The species is commonly known as the "spiny puffball"<ref name=Roody2003/> or the "spring puffball";<ref name=Dickinson1982/> Peck referred to the species as the "echinate puff-ball".<ref name=Peck1879/> The specific epithet ''echinatum'' is derived from the Greek word ''echinos'' (εχινος) meaning "hedgehog" or "sea-urchin".<ref name=Stearn2004/>
==Description== {{Multiple image|direction=vertical|align=left|image1=Lycoperdon echinatum 107913.jpg|image2=Igelstäubling Igelbovist Lycoperdon echinatum.jpg|width=175|caption2=Young specimens (above) are white before turning brown in maturity (below).}} The fruit bodies of ''L. echinatum'' are {{convert|2|–|4|cm|in|1|abbr=on}} wide by {{convert|2|–|3.5|cm|in|1|abbr=on}} tall, and are roughly spherical, or pear-shaped.<ref name=Miller2006/> The exterior surface is crowded with spines that may be up to {{convert|0.6|cm|in|1|abbr=on}} long. According to Curtis Gates Lloyd, American specimens have more slender spines than European ones.<ref name=Lloyd1905/> Initially white and becoming dark brown in maturity, the spines are often joined at the tips in groups of three or four.<ref name=Ellis1990/> In this form the puffballs resemble acorn caps of burr oak, with which they may readily be confused.<ref name=Abel1993/> The spines slough off in age, revealing a somewhat net-like or reticulated surface. The fruit body has a small base that is an off-white or purple-gray color,<ref name=Miller2006/> and it may be attached to the growing surface by thin white cords (rhizomorphs).<ref name=McKnight1987/> The internal contents of the puffball contain the gleba, a mass of spores and associated spore-producing cells. In young specimens the gleba is white and firm, but as the puffball ages, it turns yellowish and then brown to purple-brown and powdery.<ref name=Roody2003/> Mature specimens develop a pore at the top of the fruit body through which spores are released when hit by falling raindrops.<ref name=McKnight1987/>
The spores of ''L. echinatum'', roughly spherical with warts on the surface, have diameters between 4 and 6 μm.<ref name=Roody2003/> The capillitia (coarse thick-walled hyphae in the gleba) are elastic, brown in color, contain small pores, and are 5–8 μm thick.<ref name=Ellis1990/> The basidia (the spore-bearing cells) may be attached to two to four spores, and the sterigmata (tapered spine-like projections from the basidia that attach the spores) are up to 5 μm long.<ref name=Jordan2004/>
== Edibility == Like most other puffball species, ''L. echinatum'' is edible when still young and while the gleba is still white and firm.<ref name="Roody2003" /> Consumption of older specimens with a non-white gleba, or where the gleba has turned into a powdery spore mass, may cause stomach upsets.<ref name="Fischer1992" /> This species has a mild taste, and no distinguishable odor,<ref name="Miller2006" /> although one source describes the smell of dried fruit bodies as similar to "old ham".<ref>Coker ''et al''., 1974 [1928], pp. [https://books.google.com/books?id=2UrCCrPPqO4C&pg=PA73 73–5].</ref> One source notes that it is "well flavoured and tender when cooked",<ref name="Dickinson1982" /> while another describes the texture (of edible puffballs in general) as "somewhat like French toast".<ref name="Fischer1992" /> Antonio Carluccio recommends sautéeing puffballs with other mushrooms.<ref name="Carluccio2003" /> To avoid possible confusion with potentially deadly ''Amanita'' species, it is recommended to slice young puffballs with a longitudinal cut to ensure that the flesh is devoid of any internal structures.<ref name="Hall2003" />
==Similar species== [[File:Lycoperdon pulcherrimum 58681.jpg|thumb|right|''Lycoperdon pulcherrimum'' has stouter spines than ''L. echinatum''.]] ''Lycoperdon pulcherrimum'' closely resembles ''L. echinatum'', but its spines are stouter,<ref name=Miller2006/> do not turn brown in age, and the surface of the fruit body underneath the spines is smooth, not pitted.<ref name=Roody2003/> Alexander H. Smith noted that in youth, they are "difficult if not impossible to distinguish from each other, but this will cause no inconvenience to those collecting for the table, since both are edible."<ref name=Smith1951/> In some areas the two species appear to intergrade, as specimens may be found whose spines turn brown but do not fall off.<ref name=Abel1993/> Young specimens of ''L. pedicellatum'' may also be difficult to distinguish from ''L. echinatum'', but the former has a smooth outer surface when mature, and has spores attached to a pedicel (a narrow extension of the basidium on which the sterigmata and spores are formed) that is about 4–5 times as long as the spore.<ref>Coker ''et al''., 1974 [1928], pp. [https://books.google.com/books?id=2UrCCrPPqO4C&pg=PA86 85–86]. Retrieved 2010-05-17.</ref> ''Lycoperdon compactum'', found only in New Zealand, also resembles ''L. echinatum'' in appearance, but differs in having smaller spores, capillitia that are hyaline (translucent) and septate (with partitions that divide the capillitia into compartments).<ref name=Cunningham1942/>
==Habitat, distribution, and ecology== ''Lycoperdon echinatum'' can be found either solitary or in small groups. It typically grows on the ground in deciduous forests and grassy areas, glades and pastures, on moss, humus, or woody debris. The fungus has been noted to have a preference for beech woods.<ref name=Dickinson1982/><ref name=Ellis1990/> Fruit bodies may make their appearance anytime from the late spring to autumn.<ref name=Roody2003/><ref name=Miller2006/> Older specimens are more likely to be overlooked, as their brown color blends into the surrounding environment of dead leaves and dead wood.<ref name=Smith1951/> The puffball is used by various species of scuttle flies (family Phoridae) as larval food.<ref name=Disney2009/>
This species has been collected from eastern central Africa,<ref name=Demoulin1975/> China,<ref name=Bi1993/> Costa Rica,<ref name=Calonge2005/> Iran,<ref name=Saber1991/> Japan,<ref name=Kasuya2004/> and Europe (including Britain,<ref name=Holden1973/> Bulgaria,<ref name=Chalkov1985/> the Czech Republic,<ref name=Kubat1972/> Finland,<ref name=Haeggstrom1997/> Germany,<ref name=Kreisel1962/> Italy,<ref name=Belletto1977/> Slovakia,<ref name=Larsson2008/> Spain,<ref name=Ibiza1920/> Sweden,<ref name=Kers1975/><ref name=Andersson1994/> and Switzerland<ref name=Kueng1973/>). In North America, it is "locally frequent" east of the Rocky Mountains.<ref name=Dickinson1982/>
It is considered a threatened species in Åland (Finland).<ref name=Haeggstrom1997/> A study of the species' distribution in Sweden reported that in the 1940s and 50s, it grew in beech woods with broad-leaved grasses and herbs in topsoils with soil pH levels between 5.0 and 6.6, but the populations have since decreased owing to soil acidification during the last several decades.<ref name=Andersson1994/> Fruit bodies collected near arsenic-contaminated sites have been shown to bioaccumulate arsenic, largely in the form of arsenobetaine.<ref name=Slejkovec1997/>
==Antimicrobial activity== Using a standard laboratory method to determine antimicrobial susceptibility, methanol-based extracts of ''Lycoperdon umbrinum'' fruit bodies were shown in a 2005 study to have "significant" antibacterial activity against various human pathogenic bacteria, including ''Bacillus subtilis'', ''Escherichia coli'', ''Salmonella typhimurium'', ''Staphylococcus aureus'', ''Streptococcus pyogenes'', and ''Mycobacterium smegmatis''.<ref name=Dulger2005/> An earlier study (2000) had identified weak antibacterial activity against ''Enterococcus faecium'' and ''Staphylococcus aureus''.<ref name=Suay2000/> Although the specific compounds responsible for the antimicrobial activity have not been identified, chemical analysis confirms the presence of terpenoids,<ref name=Dulger2005/> a class of widely occurring organic chemicals that are being investigated for their potential use as antimicrobial drugs.<ref name=Saleem2010/>
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<ref name=Smith1951>{{cite book |title=Puffballs and their Allies in Michigan |author=Smith AH. |year=1951 |publisher=University of Michigan Press |location=Ann Arbor, Michigan |page=68 |url=http://quod.lib.umich.edu/cgi/t/text/pageviewer-idx?c=fung1tc;cc=fung1tc;q1=echinatum;rgn=full%20text;idno=agk0823.0001.001;didno=agk0823.0001.001;view=image;seq=80;page=root;size=s;frm=frameset}}</ref>
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<ref name="urlFungorum - Synonymy for Lycoperdon radicatum">{{cite web |url=http://www.speciesfungorum.org/Names/SynSpecies.asp?RecordID=163410 |title=Species Fungorum – Species synonymy for ''Lycoperdon radicatum'' |publisher=Index Fungorum. CAB International |access-date=2010-05-14}}</ref>
<ref name="urlIndex Fungorum - Synonymy for Lycoperdon gemmatum">{{cite web |url=http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=222200 |title=Species Fungorum – Species synonymy for ''Lycoperdon gemmatum''|publisher=Index Fungorum. CAB International |access-date=2010-05-17}}</ref>
<ref name="urlMycoBank: Lycoperdon hoylei">{{cite web |url=http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=172479 |title=''Lycoperdon hoylei'' Berk. & Broome 1871 |publisher=MycoBank. International Mycological Association |access-date=2010-05-14}}</ref>
<ref name="urlMycoBank: Utraria echinata">{{cite web |url=http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=197131 |title=''Utraria echinata'' (Pers.) Quél. 1873 |publisher=MycoBank. International Mycological Association |access-date=2010-05-14}}</ref>
}}
==Bibliography== *{{cite book |vauthors=Coker WS, Couch JN, Johnson MM |title=The Gasteromycetes of the Eastern United States and Canada |publisher=Dover Publications |location=New York |year=1974 |orig-year=First published 1928 |isbn=0-486-23033-3}}
==External links== {{Commons}} *{{Index Fungorum|414453}} *[http://www.grzyby.pl/gatunki/Lycoperdon_echinatum.htm Fungi of Poland] Description and several photos *[http://www.ne.jp/asahi/mushroom/tokyo/Lycoperdales/arage/arage.htm Picture of spores]
{{featured article}} {{Taxonbar|from=Q1093294}}
{{DEFAULTSORT:Lycoperdon echinatum}} echinatum Category:Edible fungi Category:Puffballs Category:Fungi of Africa Category:Fungi of Asia Category:Fungi of Central America Category:Fungi of Europe Category:Fungi of North America Category:Fungi described in 1797 Category:Taxa named by Christiaan Hendrik Persoon Category:Fungus species