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'''''Geastrum quadrifidum''''', commonly known as the '''rayed earthstar''' or '''four-footed earthstar''', is an inedible species of [[mushroom]] belonging to the genus ''[[Geastrum]]'', or earthstar fungi. First described scientifically by [[Christian Hendrik Persoon]] in 1794, ''G. quadrifidum'' is a [[cosmopolitan distribution|cosmopolitan]]—but not common—species of Europe, the Americas, Africa, Asia, and Australia. The fungus is a [[saprobe]], and spends most of its life as thin strands of [[mycelium]], feeding off decomposing organic matter present in the soil and [[forest floor|litter]] of [[coniferous forest]]s.
'''''Geastrum quadrifidum''''', commonly known as the '''rayed earthstar''' or '''four-footed earthstar''', is an inedible species of [[mushroom]] belonging to the genus ''[[Geastrum]]'', or earthstar fungi. First described scientifically by [[Christian Hendrik Persoon]] in 1794, ''G. quadrifidum'' is a [[cosmopolitan distribution|cosmopolitan]]—but not common—species of Europe, the Americas, Africa, Asia, and Australasia. The fungus is a [[saprobe]], feeding off decomposing organic matter present in the soil and [[forest floor|litter]] of [[coniferous forest]]s.


The small, tough, [[Sporocarp (fungi)|fruit bodies]] are grayish-brown balls that are initially enclosed by a skin, or [[peridium]], made of four distinct layers of tissue. The outer tissue layer splits to form star-like rays and expose a circular spore case. Inside the spore case is the ''gleba''—fertile spore-producing tissue that is white and firm when young, but becomes brown and powdery in age. The grayish-brown spore case is set on a short, slender stalk, and has a well-defined narrow pore at the top where mature spores may escape. Fully expanded, the fruit body reaches dimensions up to {{convert|2|–|3|cm|in|1|abbr=on}} wide and up to about {{convert|3|cm|in|1|abbr=on}} tall. The outer skin is purplish-brown, with four or five cream or yellowish-brown colored rays that have their tips stuck in the [[substrate (biology)|substrate]]. There is a flat weft of [[mycelium]] between ray tips. The spores are spherical, warty, and have a diameter of up to 6 [[micrometre|µm]]. ''Geastrum quadrifidum'' is one of a number of earthstars whose rays arch downward as they mature, lifting the spore sac upward, high enough to catch air currents that disseminate the spores into new habitats. The species may be easily confused with ''[[Geastrum fornicatum]]'', a larger earthstar without a well-defined pore mouth.
The small, tough, [[Sporocarp (fungi)|fruit bodies]] are grayish-brown balls that are initially enclosed by a skin, or [[peridium]], made of four distinct layers of tissue. The outer tissue layer splits to form star-like rays and expose a circular spore case. Inside the spore case is the [[gleba]]—fertile spore-producing tissue that is white and firm when young, but becomes brown and powdery in age. The grayish-brown spore case is set on a short, slender stalk, and has a well-defined narrow pore at the top where mature spores may escape. Fully expanded, the fruit body reaches dimensions up to {{convert|2|–|3|cm|in|1|abbr=on}} wide and up to about {{convert|3|cm|in|1|abbr=on}} tall. The outer skin is purplish-brown, with four or five cream or yellowish-brown colored rays that have their tips stuck in the [[substrate (biology)|substrate]]. There is a flat weft of [[mycelium]] between ray tips. The spores are spherical, warty, and have a diameter of up to 6 [[micrometre|µm]]. ''Geastrum quadrifidum'' is one of a number of earthstars whose rays arch downward as they mature, lifting the spore sac upward, high enough to catch air currents that disseminate the spores into new habitats. The species may be easily confused with ''[[Geastrum fornicatum]]'', a larger earthstar without a well-defined pore mouth.


==Taxonomy and naming==
==Taxonomy and naming==
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==Description==
==Description==
[[image:Geastrum quadrifidum spore sac closeup.JPG|thumb|left|Closeup of spore sac showing detail of peristome (above) and supporting stalk (below)]] Like all ''Geastrum'' fungi, the internal spore-producing [[gleba]] is enclosed in the [[peridium]], a protective structure composed of four layers of tissue: an inner endoperidium, and outer exoperidium that may further be divided into an external [[mycelium|mycelial]], a tough and membranous middle fibrillose layer, and an internal fleshy layer (known as the psuedoparenchyma).<ref name=Cunningham1944/> The immature, unopened fruit body is roughly spherical to somewhat flattened or irregular in shape. It lies partly or wholly submerged, encrusted with [[forest floor|debris]]. The expanded fruit body is usually taller than wide, about {{convert|10|–|40|mm|1|in|abbr=on}} high, with mycelial cup included about {{convert|15|–|55|mm|in|1|abbr=on}}. The exoperidium (the outer tissue layer of the four-layered [[peridium]]) splits in the middle into three to six, but usually four or five rays. The exoperidium is typically [[wikt:fornicate|fornicate]]—a structural feature that arises when the mesoperidium separates from the exoperidium, adhering only at the edge. In this way, the endoperidium (the internal tissue layer that encloses the spore sac) is lifted upwards with the downward movement of the rays. In this species, the tips of the rays remain attached to the mycelial layer, which remain attached to the [[substrate (biology)|substrate]] as a cup in the ground.<ref name=Sunhede1989/>
[[image:Geastrum quadrifidum spore sac closeup.JPG|thumb|left|Closeup of spore sac showing detail of peristome (above) and supporting stalk (below)]] Like all ''Geastrum'' fungi, the internal spore-producing [[gleba]] is enclosed in the [[peridium]], a protective structure composed of four layers of tissue: an inner endoperidium, and outer exoperidium that may further be divided into an external [[mycelium|mycelial]], a tough and membranous middle fibrillose layer, and an internal fleshy layer (known as the psuedoparenchyma).<ref name=Cunningham1944/> The immature, unopened fruit body is roughly spherical to somewhat flattened or irregular in shape. It lies partly or wholly submerged, encrusted with [[forest floor|debris]]. The expanded fruit body is usually taller than it is wide, about {{convert|10|–|40|mm|1|in|abbr=on}} high, with mycelial cup included about {{convert|15|–|55|mm|in|1|abbr=on}}. The exoperidium (the outer tissue layer of the four-layered [[peridium]]) splits in the middle into three to six, but usually four or five rays. The exoperidium is typically [[wikt:fornicate|fornicate]]—a structural feature that arises when the mesoperidium separates from the exoperidium, adhering only at the edge. In this way, the endoperidium (the internal tissue layer that encloses the spore sac) is lifted upwards with the downward movement of the rays. In this species, the tips of the rays remain attached to the mycelial layer, which remain attached to the [[substrate (biology)|substrate]] as a cup in the ground.<ref name=Sunhede1989/>


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===Microscopic characteristics===
===Microscopic characteristics===
The [[basidium|basidia]] of ''G.&nbsp;quadrifidum'' have a basal [[clamp connection]] or narrow into a hyphal part ending at a clamp, when young more or less ellipsoid to club-shaped, in age often becoming more or less bottle-shaped, [[wikt:ampullaceous|ampullaceous]] or sometimes almost [[wikt:lecythiform|lecythiform]] but other shapes occur, as mature (hyphal part excluded) 14–21 x 4.5–7&nbsp;[[micrometre|µm]]. The hyphal part is less than 1–6 x 1–2&nbsp;µm. The [[wikt:sterigmatum|sterigmata]] (thin projections of the basidia that attach the spore) are 4–6&nbsp;µm long, mostly 1–1.5&nbsp;µm thick. The hyphae located immediately underneath the basidia is thin-walled, 1–2&nbsp;µm wide, provided with clamps and densely branched. The hyphae of the tramal plates is roughly parallel, thin-walled, 1–2&nbsp;µm wide, and provided with clamps which may be dilated.[[image:Geastrum quadrifidum spores 84344.jpg|thumb|left|Spores are roughly spherical and have ornamentations on the surface.]]The spores in mass are dark brown when mature. They are spherical, covered with "warts" or [[wikt:verruca|verrucae]], and measure 5–6&nbsp;µm in diameter (including ornamentation).{{#tag:ref|Bates reports a slightly larger range—5.6 to 6.4&nbsp;µm—with an average of 6.1&nbsp;µm.<ref name=Bates2004/>|group=nb}} The spores often contain a drop of oil. [[Scanning electron microscopy]] reveals the verrucae to be up to 0.8&nbsp;µm long, conical to columnar processes with rounded to almost flattened tips. The [[wikt:apiculus|apiculus]] (the part of a spore which attaches to the sterigmata at the end of a basidium) is distinct with radiating ridge-like processes. The young spore is first broadly egg-shaped before becoming roughly spherical in maturity.<ref name=Sunhede1989/>
The [[basidium|basidia]] of ''G.&nbsp;quadrifidum'' have a basal [[clamp connection]] or narrow into a hyphal part ending at a clamp, when young more or less ellipsoid to club-shaped, in age often becoming more or less bottle-shaped, [[wikt:ampullaceous|ampullaceous]] or sometimes almost [[wikt:lecythiform|lecythiform]] but other shapes occur, as mature (hyphal part excluded) 14–21 x 4.5–7&nbsp;[[micrometre|µm]]. The hyphal part is less than 1–6 x 1–2&nbsp;µm. The [[wikt:sterigmatum|sterigmata]] (thin projections of the basidia that attach the spore) are 4–6&nbsp;µm long, mostly 1–1.5&nbsp;µm thick. The hyphae located immediately underneath the basidia is thin-walled, 1–2&nbsp;µm wide, provided with clamps and densely branched. The hyphae of the tramal plates is roughly parallel, thin-walled, 1–2&nbsp;µm wide, and provided with clamps which may be dilated.<ref name=Sunhede1989/>[[image:Geastrum quadrifidum spores 84344.jpg|thumb|left|Spores are roughly spherical and have ornamentations on the surface.]]The spores in mass are dark brown when mature. They are spherical, covered with "warts" or [[wikt:verruca|verrucae]], and measure 5–6&nbsp;µm in diameter (including ornamentation).{{#tag:ref|Bates reports a slightly larger range—5.6 to 6.4&nbsp;µm—with an average of 6.1&nbsp;µm.<ref name=Bates2004/>|group=nb}} The spores often contain a drop of oil. [[Scanning electron microscopy]] reveals the verrucae to be up to 0.8&nbsp;µm long, conical to columnar processes with rounded to almost flattened tips. The [[wikt:apiculus|apiculus]] (the part of a spore which attaches to the sterigmata at the end of a basidium) is distinct with radiating ridge-like processes. The young spore is first broadly egg-shaped before becoming roughly spherical in maturity.<ref name=Sunhede1989/>


The ''capillitium'' refers to coarse, late-maturing, thick-walled cells in the gleba that develop pores or slits in their thick secondary walls. The capillitial hyphae are 1.5–9.5&nbsp;µm wide, thick-walled, often with a narrow [[lumen (anatomy)|lumen]], and with or without surface ornamentation. The columella hyphae are 1.5–14&nbsp;µm wide (occasionally wider, up to 34&nbsp;µm), thick-walled, often with narrow lumen. Single thin-walled, about 1.5&nbsp;µm wide hyphae with clamps can be observed. The endoperidial hyphae are densely interwoven, thick-walled, and about 2–6&nbsp;µm wide. The whitish powder on newly expanded specimens consists of crystalline matter and thin-walled, 1.5–4&nbsp;µm wide, branched hyphae with clamps. The peristome hyphae are thick-walled, 2–11&nbsp;µm wide. The pseudoparenchymatous layer is built up by bladder-like, thin-walled hyphae of varying size. On the surface of newly expanded specimens crystals and thin-walled hyphae of the same kind as on the endoperidium are present.<ref name=Sunhede1989/> The crystals are [[calcium oxalate]] dihydrate that have the crystalline structure of a pyramid, and are arranged singly or in loose aggregates, 11 to 30&nbsp;µm in size.<ref name=Khan1995/> The fibrous layer has thick-walled hyphae 1.5–4&nbsp;µm wide. The mycelial layer in the inner, very thin part (seen as a glossy lining on the fibrous layer of newly expanded fruit bodies) consists of a dense web of thin-walled, 1.5–4&nbsp;µm wide, clamped hyphae. Thick-walled hyphae are also present, measuring 2–11 (sometimes up to 19)&nbsp;µm wide. The outer part (the mycelial cup) consists of thick-walled (often with a narrow lumen), 1.5–4&nbsp;µm wide, with branched and densely interwoven hyphae.<ref name=Sunhede1989/>
The ''capillitium'' refers to coarse, late-maturing, thick-walled cells in the gleba that develop pores or slits in their thick secondary walls. The capillitial hyphae are 1.5–9.5&nbsp;µm wide, thick-walled, often with a narrow [[lumen (anatomy)|lumen]], and with or without surface ornamentation. The columella hyphae are 1.5–14&nbsp;µm wide (occasionally wider, up to 34&nbsp;µm), thick-walled, often with narrow lumen. Single thin-walled, about 1.5&nbsp;µm wide hyphae with clamps can be observed. The endoperidial hyphae are densely interwoven, thick-walled, and about 2–6&nbsp;µm wide. The whitish powder on newly expanded specimens consists of crystalline matter and thin-walled, 1.5–4&nbsp;µm wide, branched hyphae with clamps. The peristome hyphae are thick-walled, 2–11&nbsp;µm wide. The pseudoparenchymatous layer is built up by bladder-like, thin-walled hyphae of varying size. On the surface of newly expanded specimens crystals and thin-walled hyphae of the same kind as on the endoperidium are present.<ref name=Sunhede1989/> The crystals are [[calcium oxalate]] dihydrate that have the crystalline structure of a pyramid, and are arranged singly or in loose aggregates, 11 to 30&nbsp;µm in size.<ref name=Khan1995/> The fibrous layer has thick-walled hyphae 1.5–4&nbsp;µm wide. The mycelial layer in the inner, very thin part (seen as a glossy lining on the fibrous layer of newly expanded fruit bodies) consists of a dense web of thin-walled, 1.5–4&nbsp;µm wide, clamped hyphae. Thick-walled hyphae are also present, measuring 2–11 (sometimes up to 19)&nbsp;µm wide. The outer part (the mycelial cup) consists of thick-walled (often with a narrow lumen), 1.5–4&nbsp;µm wide, with branched and densely interwoven hyphae.<ref name=Sunhede1989/>
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''Geastrum quadrifidum'' is also similar to ''[[Geastrum dissimile|G.&nbsp;dissimile]]'', ''[[Geastrum leptospermum|G.&nbsp;leptospermum]]'', and ''[[Geastrum welwitschii|G.&nbsp;welwitschii]]'' in its fruit body [[morphology (biology)|morphology]], especially the exoperidial rays, endoperidial body, and peristome. ''Geastrum dissimile'' differs from ''G.&nbsp;quadrifidum'' by its often [[wikt:sulcate|sulcate]] or silky [[wikt:fimbriate|fimbriate]], smooth peristome, and slightly smaller spores (4–5&nbsp;μm in diameter).<ref name=Bottomley1948/> ''Geastrum leptospermum'' can be distinguished from ''G.&nbsp;quadrifidum'' by its smaller spores (2–3&nbsp;μm in diameter),<ref name=Coker1974/> and by its preference for growing in mosses on tree trunks. Further, ''G.&nbsp;welwitschii'' differs from ''G.&nbsp;quadrifidum'' by its [[epigeal]] mycelial cup with a felted or tufted outer surface, and indistinctly delimited peristome.<ref name=Sunhede1989/>
''Geastrum quadrifidum'' is also similar to ''[[Geastrum dissimile|G.&nbsp;dissimile]]'', ''[[Geastrum leptospermum|G.&nbsp;leptospermum]]'', and ''[[Geastrum welwitschii|G.&nbsp;welwitschii]]'' in its fruit body [[morphology (biology)|morphology]], especially the exoperidial rays, endoperidial body, and peristome. ''Geastrum dissimile'' differs from ''G.&nbsp;quadrifidum'' by its often [[wikt:sulcate|sulcate]] or silky [[wikt:fimbriate|fimbriate]], smooth peristome, and slightly smaller spores (4–5&nbsp;μm in diameter).<ref name=Bottomley1948/> ''Geastrum leptospermum'' can be distinguished from ''G.&nbsp;quadrifidum'' by its smaller spores (2–3&nbsp;μm in diameter),<ref name=Coker1974/> and by its preference for growing in mosses on tree trunks. Further, ''G.&nbsp;welwitschii'' differs from ''G.&nbsp;quadrifidum'' by its [[epigeal]] mycelial cup with a felted or tufted outer surface, and indistinctly delimited peristome.<ref name=Sunhede1989/>


==Distribution and habitat==
==Distribution, ecology, and habitat==
''Geastrum quadrifidum'' has a widespread distribution. European countries from which the fungus has been reported include Belgium,<ref name=Demoulin1968/> France,<ref name=Anon2006/> Germany,<ref name=Winterhoff1981/> Norway,<ref name=Hapnes1990/> Poland,<ref name=Wojewoda2000/> and Sweden.<ref name=Andersson2010/> In Asia, it has been collected from China and Japan.<ref name=Kasuya2009/> The North American distribution extends from Canada<ref name=Schalkwijk-Barendsen1991/> south to Mexico,<ref name=Esqueda2003/> and includes Hawaii.<ref name=Smith1982/> It is also found in Australia<ref name=May2003/> and New Zealand, South Africa, and South America.<ref name=PoncedeLeon1968/>
Although ''Geastrum quadrifidum'' has a widespread distribution, it is not a common species. European countries from which the fungus has been reported include Belgium,<ref name=Demoulin1968/> France,<ref name=Anon2006/> Germany,<ref name=Winterhoff1981/> Norway,<ref name=Hapnes1990/> Poland,<ref name=Wojewoda2000/> and Sweden.<ref name=Andersson2010/> In Asia, it has been collected from China and Japan.<ref name=Kasuya2009/> The North American distribution extends from Canada<ref name=Schalkwijk-Barendsen1991/> south to Mexico,<ref name=Esqueda2003/> and includes Hawaii.<ref name=Smith1982/> It is also found in Australia<ref name=May2003/> and New Zealand, South Africa, and South America.<ref name=PoncedeLeon1968/> Because of its rarity, it has been placed on the [[Regional Red List]]s of several European countries, including [[Montenegro]],<ref name=Peric2005/> Denmark,<ref name="urlNERI - The Danish Red Data Book - Geastrum quadrifidum Pers.: Pers."/> Norway,<ref name="urlRED LIST NORWAY"/> and Poland.<ref name=Wojewoda2000/>


⚫
Like most earthstars, ''G.&nbsp;quadrifidum'' is a [[saprobic]] fungus, and spends most of its [[biological life cycle|life cycle]] as thin strands of [[mycelium]], deriving nutrients by decomposing [[forest floor|leaf litter]] and similar [[detritus]], converting it to [[humus]] and [[mineralization|mineralizing]] organic matter in the soil.<ref name=Miller1988/> The fungal fruit bodies are generally found in [[coniferous forest|coniferous woodland]], where they appear in summer and autumn. In Mexico, it was found in tropical [[thorn forest]] and [[Sierra Madre Oriental pine-oak forests|pine-oak forest]] in the summer.<ref name=Esqueda2003/> In Britain, all collections have been made in [[beech]] forest on [[calcareous]] soil.<ref name=Pegler1995/>
Because of its rarity, it has been placed on the [[Regional Red List]]s of several European countries, including [[Montenegro]],<ref name=Peric2005/> Denmark,<ref name="urlNERI - The Danish Red Data Book - Geastrum quadrifidum Pers.: Pers."/> Norway,<ref name="urlRED LIST NORWAY"/> and Poland.<ref name=Wojewoda2000/>

⚫
The fungus is generally found in [[coniferous forest|coniferous woodland]], where it fruits in the summer and autumn. In Mexico, it was found in tropical [[thorn forest]] and [[Sierra Madre Oriental pine-oak forests|pine-oak forest]] in the summer.<ref name=Esqueda2003/> In Britain, all collections have been made in [[beech]] forest on [[calcareous]] soil.<ref name=Pegler1995/>


==Notes==
==Notes==
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<ref name=Andersson2010>{{cite journal |author=Andersson U-B. |year=2010 |title=Jordstjärnor i Sverige 5. Fyrflikig jordstjärna, hög jordstjärna, hårig jordstjärna, sålljordstjärna |trans_title=Swedish earthstars (Geastraceae) 5 |journal=Svensk Botanisk Tidskrift |volume=104 |issue=1 |pages=39–43 |language=Swedish}}</ref>
<ref name=Andersson2010>{{cite journal |author=Andersson U-B. |year=2010 |title=Jordstjärnor i Sverige 5. Fyrflikig jordstjärna, hög jordstjärna, hårig jordstjärna, sålljordstjärna |trans_title=Swedish earthstars (Geastraceae) 5 |journal=Svensk Botanisk Tidskrift |volume=104 |issue=1 |pages=39–43 |language=Swedish}}</ref>


<ref name=Anon2006>{{cite journal |author=Anonymous. |year=2006 |trans_title=Top of the basket 2 |journal=Bulletin de la Société Linnéenne de Bordeaux |volume=34 |issue=4 |pages=307–09 |language=French}}</ref>
<ref name=Anon2006>{{cite journal |author=Anonymous. |year=2006 |title=Le dessus du panier n° 2 |trans_title=Top of the basket 2 |journal=Bulletin de la Société Linnéenne de Bordeaux |volume=34 |issue=4 |pages=307–09 |language=French}}</ref>


<ref name=Bates2004>{{Cite thesis |degree=M.Sc. |chapter=Taxonomy |title=Arizona Members of the Geastraceae and Lycoperdaceae (Basidiomycota, Fungi) |url=http://www.azfungi.org/stbates/STBThesis2.pdf |format=PDF|author=Bates ST. |year=2004 |publisher=Arizona State University |accessdate=2010-06-20 |pages=119–22}}</ref>
<ref name=Bates2004>{{Cite thesis |degree=M.Sc. |chapter=Taxonomy |title=Arizona Members of the Geastraceae and Lycoperdaceae (Basidiomycota, Fungi) |url=http://www.azfungi.org/stbates/STBThesis2.pdf |format=PDF|author=Bates ST. |year=2004 |publisher=Arizona State University |accessdate=2010-06-20 |pages=119–22}}</ref>
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<ref name=May2003>{{cite book |title=Fungi of Australia. Basidiomycota p.p. & Myxomycota |volume=2B |last=May TW, Milne J, Shingles S, Jones RH. |year=2008 |publisher=CSIRO Publishing |location= |isbn=978-0643069077 |page=117 |url=http://books.google.com/books?id=mTJsa5a3ZpcC&lpg=PA117&dq=Geastrum%20quadrifidum&pg=PA117#v=onepage&q=Geastrum%20quadrifidum&f=false |accessdate=2010-06-20}}</ref>
<ref name=May2003>{{cite book |title=Fungi of Australia. Basidiomycota p.p. & Myxomycota |volume=2B |last=May TW, Milne J, Shingles S, Jones RH. |year=2008 |publisher=CSIRO Publishing |location= |isbn=978-0643069077 |page=117 |url=http://books.google.com/books?id=mTJsa5a3ZpcC&lpg=PA117&dq=Geastrum%20quadrifidum&pg=PA117#v=onepage&q=Geastrum%20quadrifidum&f=false |accessdate=2010-06-20}}</ref>

<ref name=Miller1988>{{cite book |author=Miller HR, Miller OK. |title=Gasteromycetes: Morphological and Developmental Features, with Keys to the Orders, Families, and Genera |publisher=Mad River Press |location=Eureka, CA |year=1988 |pages=31, 36 |isbn=0-916422-74-7}}</ref>


<ref name=Pegler1995>{{cite book |author=Læssøe T, Pegler DN, Spooner B. |title=British Puffballs, Earthstars and Stinkhorns: an Account of the British Gasteroid Fungi |publisher=Royal Botanic Gardens |location=Kew |year=1995 |page=98 |isbn=0-947643-81-8}}</ref>
<ref name=Pegler1995>{{cite book |author=Læssøe T, Pegler DN, Spooner B. |title=British Puffballs, Earthstars and Stinkhorns: an Account of the British Gasteroid Fungi |publisher=Royal Botanic Gardens |location=Kew |year=1995 |page=98 |isbn=0-947643-81-8}}</ref>
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<ref name=Wojewoda2000>{{cite journal |author=Wojewoda W. |year=2000 |title=New localities of rare and threatened species of Geastrum (Lycoperdales) in Poland |journal=Acta Mycologica |volume=35 |issue=2 |pages=145–51 |issn=0001-625X}}</ref>
<ref name=Wojewoda2000>{{cite journal |author=Wojewoda W. |year=2000 |title=New localities of rare and threatened species of Geastrum (Lycoperdales) in Poland |journal=Acta Mycologica |volume=35 |issue=2 |pages=145–51 |issn=0001-625X}}</ref>

<!--



<ref name=Dissing1962>{{cite journal |author= Dissing H, Lange M|year= 1962 |title=Gasteromycetes of Congo |journal=Bulletin du Jardin botanique de l'État |volume=32 |issue=4 |pages=325–416 |doi=10.2307/3667249}}</ref>

<ref name=Ellis1990>{{cite book |author=Ellis JB, Ellis MB|title=Fungi without Gills (Hymenomycetes and Gasteromycetes): an Identification Handbook |publisher=Chapman and Hall |location=London |year=1990 |pages=226–27 |isbn=0-412-36970-2}}</ref>

<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 |pages=86–87 |oclc= 1002833}}</ref>

{{cite journal |author= |year= |title= |journal= |volume= |issue= |pages= |url= |doi= |pmid= |pmc= }}

-->


}}
}}
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[[Category:Geastrum|quadrifidum]]
[[Category:Geastrum|quadrifidum]]
[[Category:Fungi of Asia]]
[[Category:Fungi of Asia]]
[[Category:Fungi of Africa]]
[[Category:Fungi of Australia]]
[[Category:Fungi of Australia]]
[[Category:Fungi of Europe]]
[[Category:Fungi of Europe]]
[[Category:Fungi of New Zealand]]
[[Category:Fungi of New Zealand]]
[[Category:Fungi of North America]]
[[Category:Fungi of North America]]
[[Category:Fungi of South America]]
[[Category:Inedible fungi]]
[[Category:Inedible fungi]]
{{Good article}}
{{Good article}}

Revision as of 03:15, 25 September 2010

Template:FixBunching

Geastrum quadrifidum
Scientific classification
Kingdom:
Fungi
Phylum:
Basidiomycota
Class:
Agaricomycetes
Order:
Geastrales
Family:
Geastraceae
Genus:
Geastrum
Species:
G. quadrifidum
Binomial name
Geastrum quadrifidum
Pers.:Pers.
Synonyms[2]

Lycoperdon coronatum Schaeff.
Lycoperdon coronatum Scop.
Geaster coronatus (Schaeff.) J. Schröt.
Geastrum quadrifidum var. minus Pers.
Geastrum minus (Pers.) G. Cunn.[1] auct. non

Template:FixBunching

Geastrum quadrifidum
View the Mycomorphbox template that generates the following list
Glebal hymenium
No distinct cap
Spore print is brown
Ecology is saprotrophic
Edibility is inedible

Template:FixBunching Geastrum quadrifidum, commonly known as the rayed earthstar or four-footed earthstar, is an inedible species of mushroom belonging to the genus Geastrum, or earthstar fungi. First described scientifically by Christian Hendrik Persoon in 1794, G. quadrifidum is a cosmopolitan—but not common—species of Europe, the Americas, Africa, Asia, and Australasia. The fungus is a saprobe, feeding off decomposing organic matter present in the soil and litter of coniferous forests.

The small, tough, fruit bodies are grayish-brown balls that are initially enclosed by a skin, or peridium, made of four distinct layers of tissue. The outer tissue layer splits to form star-like rays and expose a circular spore case. Inside the spore case is the gleba—fertile spore-producing tissue that is white and firm when young, but becomes brown and powdery in age. The grayish-brown spore case is set on a short, slender stalk, and has a well-defined narrow pore at the top where mature spores may escape. Fully expanded, the fruit body reaches dimensions up to 2–3 cm (0.8–1.2 in) wide and up to about 3 cm (1.2 in) tall. The outer skin is purplish-brown, with four or five cream or yellowish-brown colored rays that have their tips stuck in the substrate. There is a flat weft of mycelium between ray tips. The spores are spherical, warty, and have a diameter of up to 6 µm. Geastrum quadrifidum is one of a number of earthstars whose rays arch downward as they mature, lifting the spore sac upward, high enough to catch air currents that disseminate the spores into new habitats. The species may be easily confused with Geastrum fornicatum, a larger earthstar without a well-defined pore mouth.

Taxonomy and naming

The Dutch mycologist Christian Hendrik Persoon published the first official description of Geastrum quadrifidum in 1794,[3] and later sanctioned this name in his 1801 Synopsis Methodica Fungorum (as Geastrum quadrifidum var. minus, a variety now considered synonymous with G. quadrifidum).[4] Although the species had been previously described as Lycoperdon coronatum by Jacob Christian Schaeffer (1763) and Giovanni Antonio Scopoli (1772),[5] then afterward as Geaster coronatus by Joseph Schröter (1889),[6][nb 1] the epithet coronatus is not to be used because of the existence of the sanctioned name.[8]

In Japan, G. quadrifidum has occasionally been collected under the name "Geastrum minus" (Pers.) G. Cunn. (for example, as in Imai, 1936);[9][10] within taxonomical terminology, this usage is an auctorum non—a misapplication or misinterpretation of the species name.

According to Stanek's classification of the genus Geastrum,[11] (a classification later endorsed in Sunhede's 1989 monograph of European Geastrum species), G. quadrifidum belongs in the subsection Glabrostoma of the section Perimyceliata, a grouping of similar Geastrum species that incorporate and encrust debris in the mycelial layer, and have an even peristome (opening) that is fibrillose (made of more or less parallel thin thread-like filaments). Several common names for G. quadrifidum have been suggested, including the "rayed earthstar",[12] the "four-pointed earthstar"[13] or the "four-footed earthstar".[14] Samuel Frederick Gray called it the "four-cut shell-puff" in his 1821 The Natural Arrangement of British Plants,[15] but the name was not adopted by future authors. The specific epithet quadrifidum is derived from Latin, and means "four-forks".[16]

Description

Closeup of spore sac showing detail of peristome (above) and supporting stalk (below)

Like all Geastrum fungi, the internal spore-producing gleba is enclosed in the peridium, a protective structure composed of four layers of tissue: an inner endoperidium, and outer exoperidium that may further be divided into an external mycelial, a tough and membranous middle fibrillose layer, and an internal fleshy layer (known as the psuedoparenchyma).[17] The immature, unopened fruit body is roughly spherical to somewhat flattened or irregular in shape. It lies partly or wholly submerged, encrusted with debris. The expanded fruit body is usually taller than it is wide, about 10–40 mm (0.4–1.6 in)* high, with mycelial cup included about 15–55 mm (0.6–2.2 in). The exoperidium (the outer tissue layer of the four-layered peridium) splits in the middle into three to six, but usually four or five rays. The exoperidium is typically fornicate—a structural feature that arises when the mesoperidium separates from the exoperidium, adhering only at the edge. In this way, the endoperidium (the internal tissue layer that encloses the spore sac) is lifted upwards with the downward movement of the rays. In this species, the tips of the rays remain attached to the mycelial layer, which remain attached to the substrate as a cup in the ground.[18]

Fresh fruit bodies have a fleshy white to cream-colored exoperidium. This individual has a pseudoparenchymatous collar, the thickness of which hides the short stalk that supports the spore sac.
When the pseudoparenchymatous layer dries it shrinks and hardens, darkening to brown; the structure is maintained by the underlying stiff papery fibrous layer.

Unlike some other Geastrum species, the rays of G. quadrifidum are not hygroscopic: they do not open and close in response to changes in humidity. Generally, the rays are broad, but may seem narrow as their edges are often rolled inwards. The width of the exoperidium (when still attached to the mycelial cup) is 8–25 mm (0.3–1.0 in)*, when fully expanded about 15–90 mm (0.6–3.5 in)*. The pseudoparenchymatous layer when fresh is about 1–2 mm thick, initially whitish, later turning beige to brownish (sometimes over reddish tints), and dark brown when old. In newly expanded specimens this layer is covered with a thin layer of crystals and hyphae, sometimes forming a pseudoparenchymatous cup or collar that often peels off in patches, when dry shrunken and hard. The fibrous layer is papery to leathery. The inner side, when free from pseudoparenchymatous remnants, is almost white, in age becoming dirty grayish-white and sometimes greenish due to algae; the outer side is initially whitish, somewhat glossy, but in age becomes grayish-white and dull. The mycelial layer has a whitish inner side and is strongly attached to the litter on its outer side. It persists for a long time (1–2-year-old fruit bodies with intact mycelial cups have been found).[18]

The spore sac is variable in shape, ranging from roughly spherical to somewhat egg-shaped, to more or less irregularly shaped, but it is usually taller than it is wide. Its diameter ranges between 3.5–16 mm (0.14–0.63 in), although it is most commonly between 5–10 mm (0.2–0.4 in). An apophysis (a swelling on the underside of the spore sac) is often present. The stalk is visible when the pseudoparenchymatous layer has dried up, and is short but distinct, measuring 1–2.5 mm tall. The color is variable; in dry specimens it is whitish, light beige, beige gray, smokey gray or brownish-gray. The endoperidium in newly expanded fruit bodies is pruinose: covered with a light beige to whitish powder of hyphae and crystalline matter. This powder gradually disappears as the fruit body ages. Its color is highly variable, and both light and dark endoperidia are present. The peristome (a clearly demarcated region encircling the opening of the spore sac) is distinctly delimited, with a disc-like to more or less conical shape.[18] It is lighter in color than the spore sac, and up to 2 mm high.[19] In old specimens, the hyphae around the peristome sometimes sticks together to develop radial grooves. The color is variable, but often of grayish or grayish-brown tints, often lighter than the endoperidium. The columella (sterile tissue, usually originating in the base of the gleba, extending into or through the gleba) is rather weak, more or less columnar to club-shaped, emerging from a more or less bulge-like continuation of the stalk and intruding to about the half or more into the mature gleba. The mature gleba is dark brown.[18] G. quadrifidum is inedible.[16]

Microscopic characteristics

The basidia of G. quadrifidum have a basal clamp connection or narrow into a hyphal part ending at a clamp, when young more or less ellipsoid to club-shaped, in age often becoming more or less bottle-shaped, ampullaceous or sometimes almost lecythiform but other shapes occur, as mature (hyphal part excluded) 14–21 x 4.5–7 µm. The hyphal part is less than 1–6 x 1–2 µm. The sterigmata (thin projections of the basidia that attach the spore) are 4–6 µm long, mostly 1–1.5 µm thick. The hyphae located immediately underneath the basidia is thin-walled, 1–2 µm wide, provided with clamps and densely branched. The hyphae of the tramal plates is roughly parallel, thin-walled, 1–2 µm wide, and provided with clamps which may be dilated.[18]

Spores are roughly spherical and have ornamentations on the surface.

The spores in mass are dark brown when mature. They are spherical, covered with "warts" or verrucae, and measure 5–6 µm in diameter (including ornamentation).[nb 2] The spores often contain a drop of oil. Scanning electron microscopy reveals the verrucae to be up to 0.8 µm long, conical to columnar processes with rounded to almost flattened tips. The apiculus (the part of a spore which attaches to the sterigmata at the end of a basidium) is distinct with radiating ridge-like processes. The young spore is first broadly egg-shaped before becoming roughly spherical in maturity.[18]

The capillitium refers to coarse, late-maturing, thick-walled cells in the gleba that develop pores or slits in their thick secondary walls. The capillitial hyphae are 1.5–9.5 µm wide, thick-walled, often with a narrow lumen, and with or without surface ornamentation. The columella hyphae are 1.5–14 µm wide (occasionally wider, up to 34 µm), thick-walled, often with narrow lumen. Single thin-walled, about 1.5 µm wide hyphae with clamps can be observed. The endoperidial hyphae are densely interwoven, thick-walled, and about 2–6 µm wide. The whitish powder on newly expanded specimens consists of crystalline matter and thin-walled, 1.5–4 µm wide, branched hyphae with clamps. The peristome hyphae are thick-walled, 2–11 µm wide. The pseudoparenchymatous layer is built up by bladder-like, thin-walled hyphae of varying size. On the surface of newly expanded specimens crystals and thin-walled hyphae of the same kind as on the endoperidium are present.[18] The crystals are calcium oxalate dihydrate that have the crystalline structure of a pyramid, and are arranged singly or in loose aggregates, 11 to 30 µm in size.[20] The fibrous layer has thick-walled hyphae 1.5–4 µm wide. The mycelial layer in the inner, very thin part (seen as a glossy lining on the fibrous layer of newly expanded fruit bodies) consists of a dense web of thin-walled, 1.5–4 µm wide, clamped hyphae. Thick-walled hyphae are also present, measuring 2–11 (sometimes up to 19) µm wide. The outer part (the mycelial cup) consists of thick-walled (often with a narrow lumen), 1.5–4 µm wide, with branched and densely interwoven hyphae.[18]

Similar species

Geastrum quadrifidum may be readily confused with G. fornicatum, which is larger—up to 15 cm (5.9 in)—and has smaller spores (4–5 μm in diameter).[21] Geastrum minimum, although similarly diminutive, is distinguished by having more rays (usually more than seven), and it is not fornicate.[22] Also, its mycelial layer is attached to the fibrous layer for a long time, without forming a mycelial cup like G. quadrifidum.[10] The Chilean species G. jurei does not have a clearly demarcated peristome.[23]

Geastrum quadrifidum is also similar to G. dissimile, G. leptospermum, and G. welwitschii in its fruit body morphology, especially the exoperidial rays, endoperidial body, and peristome. Geastrum dissimile differs from G. quadrifidum by its often sulcate or silky fimbriate, smooth peristome, and slightly smaller spores (4–5 μm in diameter).[24] Geastrum leptospermum can be distinguished from G. quadrifidum by its smaller spores (2–3 μm in diameter),[25] and by its preference for growing in mosses on tree trunks. Further, G. welwitschii differs from G. quadrifidum by its epigeal mycelial cup with a felted or tufted outer surface, and indistinctly delimited peristome.[18]

Distribution, ecology, and habitat

Although Geastrum quadrifidum has a widespread distribution, it is not a common species. European countries from which the fungus has been reported include Belgium,[22] France,[26] Germany,[27] Norway,[28] Poland,[29] and Sweden.[30] In Asia, it has been collected from China and Japan.[10] The North American distribution extends from Canada[13] south to Mexico,[31] and includes Hawaii.[32] It is also found in Australia[33] and New Zealand, South Africa, and South America.[34] Because of its rarity, it has been placed on the Regional Red Lists of several European countries, including Montenegro,[35] Denmark,[36] Norway,[37] and Poland.[29]

Like most earthstars, G. quadrifidum is a saprobic fungus, and spends most of its life cycle as thin strands of mycelium, deriving nutrients by decomposing leaf litter and similar detritus, converting it to humus and mineralizing organic matter in the soil.[38] The fungal fruit bodies are generally found in coniferous woodland, where they appear in summer and autumn. In Mexico, it was found in tropical thorn forest and pine-oak forest in the summer.[31] In Britain, all collections have been made in beech forest on calcareous soil.[39]

Notes

  1. ^ This is not to be confused with the similarly named Geastrum coronatum, a valid and distinct species that is independent of G. quadrifidum.[7]
  2. ^ Bates reports a slightly larger range—5.6 to 6.4 µm—with an average of 6.1 µm.[19]

References

  1. ^ Cunningham GH. (1926). "The Gasteromycetes of Australasia. IV. Species of the genus Geaster". The Proceedings of the Linnean Society of New South Wales. 51 (206, part 2): 72–93.
  2. ^ "Geastrum quadrifidum Pers. 1794". MycoBank. International Mycological Association. Retrieved 2010-06-20.
  3. ^ Persoon CH. (1794). "Dispositio methodica fungorum". Neues Magazin für die Botanik, Römer (in Latin). 1: 86. {{cite journal}}: Cite has empty unknown parameter: |1= (help)
  4. ^ Persoon CH. (1801). Synopsis Methodica Fungorum. 1. Göttingen: 133 http://194.203.77.76/librifungorum/Image.asp?ItemID=81&ImageFileName=0133b.jpg. Retrieved 2010-06-20. {{cite journal}}: Missing or empty |title= (help)
  5. ^ Scopoli JA. (1771). Flora carniolica (2 ed.). {{cite book}}: Unknown parameter |volme= ignored (help)
  6. ^ Schröter J. (1889). Kryptogamen-Flora von Schlesien (in German). Vol. 3–1(6). Breslau: J.U. Kern's Verlag. p. 702. {{cite book}}: Unknown parameter |trans_title= ignored (|trans-title= suggested) (help)
  7. ^ "Geastrum coronatum Pers. 1801". MycoBank. International Mycological Association. Retrieved 2010-06-13.
  8. ^ International Code of Botanical Nomenclature (Vienna Code, 2005) Article 15.
  9. ^ Imai S. (1936). "Symbolae ad floram mycologicam Asiae Orientalis. I". Botanical Magazine (Tokyo). 50 (592): 216–24. Retrieved 2010-05-20.
  10. ^ a b c Kasuya T, Yamamoto Y, Sakamoto H, Takehashi S, Hoshiino T, Kobayashi T. (2009). "Floristic study of Geastrum in Japan: three new records for Japanese mycobiota and reexamination of the authentic specimen of Geastrum minus reported by Sanshi Imai". Mycoscience. 50 (2): 84–93. doi:10.1007/s10267-008-0461-1.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  11. ^ Published in: Pilát A (1958). Gasteromycetes, Houby-Břichatky. Flora ČSR B1 (in Czech). Prague, Czechoslovakia: Nakladatelstvi Československé Akademie Vĕd. {{cite book}}: Unknown parameter |trans_title= ignored (|trans-title= suggested) (help)
  12. ^ Phillips R. "Geastrum quadrifidum". Rogers Mushrooms. Retrieved 2010-06-14.
  13. ^ a b Schalkwijk-Barendsen HME. (1991). Mushrooms of Western Canada. Edmonton, Canada: Lone Pine Publishing. p. 349. ISBN 0-919433-47-2.
  14. ^ "Recommended English Names for Fungi in the UK" (PDF) (PDF). British Mycological Society. Retrieved 2010-06-20.
  15. ^ Gray SF. (1821). A Natural Arrangement of British Plants. London: Baldwin, Cradock, and Joy, Pasternoster-Row. p. 585. Retrieved 2010-06-20.
  16. ^ a b Evenson VS. (1997). Mushrooms of Colorado and the Southern Rocky Mountains. Westcliffe Publishers. p. 189. ISBN 978-1565791923. Retrieved 2010-06-20.
  17. ^ Cunningham GH (1944). The Gasteromycetes of Australia and New Zealand. Dunedin: McIndoe. pp. 160–61.
  18. ^ a b c d e f g h i Sunhede, 1989, p. 338.
  19. ^ a b Bates ST. (2004). "Taxonomy". Arizona Members of the Geastraceae and Lycoperdaceae (Basidiomycota, Fungi) (PDF) (M.Sc. thesis). Arizona State University. pp. 119–22. Retrieved 2010-06-20.
  20. ^ Khan SR. (1995). Calcium Oxalate in Biological Systems. Boca Raton: CRC Press. p. 72. ISBN 0-8493-7673-4. Retrieved 2010-06-20.
  21. ^ Sunhede, 1989, p. 209.
  22. ^ a b Demoulin V. (1968). "Gastéromycètes de Belgique: Sclerodermatales, Tulostamatales, Lycoperdales". Bulletin du Jardin Botanique National de Belgique (in French). 38: 1–101. doi:10.2307/3667475. {{cite journal}}: Unknown parameter |trans_title= ignored (|trans-title= suggested) (help)
  23. ^ Lazo W. (1972). "Fungi from Chile I. Some Gasteromycetes and Agaricales". Mycologia. 64 (4): 786–98. JSTOR 3757933.
  24. ^ Bottomley AM. (1948). "Gasteromycetes of South Africa". Bothalia. 4: 473–810.
  25. ^ Johnson MM, Coker WS, Couch JN. (1974) [First published 1928]. The Gasteromycetes of the Eastern United States and Canada. New York: Dover Publications. ISBN 0-486-23033-3.{{cite book}}: CS1 maint: multiple names: authors list (link)
  26. ^ Anonymous. (2006). "Le dessus du panier n° 2". Bulletin de la Société Linnéenne de Bordeaux (in French). 34 (4): 307–09. {{cite journal}}: Unknown parameter |trans_title= ignored (|trans-title= suggested) (help)
  27. ^ Winterhoff W. (1981). "Alte und neue Erdsternfunde im Flugsandgebiet zwischen Walldorf und Mainz". Hessische Floristische Briefe (in German). 30 (2): 18–27. ISSN 0439-0687. {{cite journal}}: Unknown parameter |trans_title= ignored (|trans-title= suggested) (help)
  28. ^ Hapnes A, Often A. (1990). "Geastrum on Islands in Snasa Lake Nord-Trondelag Norway". Blyttia (in Norwegian). 48 (4): 155–56. ISSN 0006-5269.
  29. ^ a b Wojewoda W. (2000). "New localities of rare and threatened species of Geastrum (Lycoperdales) in Poland". Acta Mycologica. 35 (2): 145–51. ISSN 0001-625X.
  30. ^ Andersson U-B. (2010). "Jordstjärnor i Sverige 5. Fyrflikig jordstjärna, hög jordstjärna, hårig jordstjärna, sålljordstjärna". Svensk Botanisk Tidskrift (in Swedish). 104 (1): 39–43. {{cite journal}}: Unknown parameter |trans_title= ignored (|trans-title= suggested) (help)
  31. ^ a b Esqueda M, Herrera T, Perez-Siva E, Sanchez A. (2003). "Distribution of Geastrum species from some priority regions for conservation of biodiversity of Sonora, Mexico". Mycotaxon. 87: 445–56. Retrieved 2010-06-07.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  32. ^ Smith CW, Ponce de Léon P. (1982). "Hawaiian geastroid fungi". Mycologia. 74 (5): 712–17. doi:10.2307/3792856.
  33. ^ May TW, Milne J, Shingles S, Jones RH. (2008). Fungi of Australia. Basidiomycota p.p. & Myxomycota. Vol. 2B. CSIRO Publishing. p. 117. ISBN 978-0643069077. Retrieved 2010-06-20.{{cite book}}: CS1 maint: multiple names: authors list (link)
  34. ^ Ponce de Leon P. (1968). "A Revision of the Family Geastraceae". Fieldania: Botany. 31: 303–52.
  35. ^ Peric B, Peric O. (2005). The provisory red list of endangered macromycets of Montenegro (PDF) (Report). Eidg. Forschungsanstalt für Wald, Schnee und Landschaft WSL (Swiss Federal Institute for Forest, Snow and Landscape Research). Retrieved 2010-06-20.
  36. ^ "NERI - The Danish Red Data Book - Geastrum quadrifidum Pers.: Pers". Danmarks Miljøundersøgelser: National Environmental Research Institute, Denmark. Retrieved 2010-06-14.
  37. ^ "RED LIST OF THREATENED FUNGI IN NORWAY". Norsk Rødliste 2006. The Herbarium, The Natural History Museums and Botanical Garden, University of Oslo. Retrieved 2010-06-14.
  38. ^ Miller HR, Miller OK. (1988). Gasteromycetes: Morphological and Developmental Features, with Keys to the Orders, Families, and Genera. Eureka, CA: Mad River Press. pp. 31, 36. ISBN 0-916422-74-7.
  39. ^ Læssøe T, Pegler DN, Spooner B. (1995). British Puffballs, Earthstars and Stinkhorns: an Account of the British Gasteroid Fungi. Kew: Royal Botanic Gardens. p. 98. ISBN 0-947643-81-8.{{cite book}}: CS1 maint: multiple names: authors list (link)

Cited books

  • Sunhede S. (1989). Geastraceae (Basidiomycotina): Morphology, Ecology, and Systematics with Special Emphasis on the North European Species. Synopsis Fungorum, 1. Oslo, Norway: Fungiflora. ISBN 82-90724-05-5.

External links

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