Geastrum pectinatum

Geastrum pectinatum is an inedible species of mushroom belonging to the earthstar family of fungi. Although young specimens are spherical, fruit body development involves the outer layer of tissue splitting open like a star into 7 to 10 pointed rays that eventually bend back to point downward, revealing a small – 1 to 2.5 cm (0.4 to 1.0 in) broad – spore sac. The spore sac is supported by a small radially wrinkled stalk. There is a distinct conical opening (peristome) at the top of the spore sac that is up to 8 mm (0.3 in) long. It is commonly known as the beaked earthstar or the beret earthstar, in reference to the shape of the spore sac and its prominent, protruding peristome. The mass of spores and surrounding cells within the sac, the gleba, is dark-brown, and becomes powdery in mature specimens. Spores are spherical, measuring 4 to 6 micrometers in diameter, with warts on their surfaces. Although uncommon, Geastrum pectinatum has a cosmopolitan distribution, and has been collected in various locations in Europe, North and South America, Asia, Australia and Africa, where it grows on the ground in open woods. Like several other earthstars, crystals of calcium oxalate are found on G. pectinatum, and are thought to be involved in fruit body maturation.

Geastrum pectinatum
Scientific classification
Kingdom:
Fungi
Division:
Basidiomycota
Class:
Agaricomycetes
Order:
Geastrales
Family:
Geastraceae
Genus:
Geastrum
Species:
G. pectinatum
Binomial name
Geastrum pectinatum
Pers. (1801)
Synonyms[1][2]

G. plicatum Berk. (1839)
G. tenuipes Berk. (1848)
G. biplicatum Berk. & M.A.Curtis (1858)
G. pectinatum var. tenuipes (Berk.) Cleland & Cheel (1915)

Geastrum pectinatum
float
Mycological characteristics
glebal hymenium
no distinct cap
spore print is brown
ecology is saprotrophic
edibility: inedible

Taxonomy, classification, and naming

Christian Hendrik Persoon published the first description of Geastrum pectinatum in 1801.[3] In 1860, Miles Joseph Berkeley and Moses Ashley Curtis described the species Geastrum biplicatum (originally named Geaster biplicatus),[4] based on specimens sent to them by Charles Wright that he obtained from the Bonin Islands during the North Pacific Exploring and Surveying Expedition. Japanese mycologist Sanshi Imai considered this identical with G. pectinatum in a 1936 publication.[2] In 1959, mycologist J.T. Palmer reported comparing the original specimen collected by Persoon with fresh samples of what were then thought to be the distinct species G. plicatum and G. tenuipes (named by English naturalist Miles Joseph Berkeley in 1838[5] and 1848,[6] respectively) and concluded the three specimens were synonymous; the original Persoon specimen was then designated as the neotype.[7]

In Ponce de Leon's classification of Geastrum, he placed the species in the subgenus Geastrum, section Geastrum, as the type of the subsection Sulcostomata, group Pectinatum. Other species in this group—characterized by a determinate peristome surrounded by a groove—are G. xerophilum, and G. furfuraceum.[8] In Stanek's (1958) infrageneric concept, G. pectinatum is placed in section Perimyceliata (encompassing species whereby the mycelial layer covers the entire endoperidium), in subsection Glabrostomata, which includes species with plicate peristomes.[9]

The specific epithet is derived from the Latin pectinatum, "like a comb".[10] Its common names include the "beaked earthstar" or the "beret earthstar".[10] Samuel Frederick Gray called it the "comblike shell-puff" in his 1821 "A Natural Arrangement of British Plants".[11]

Description

Immature specimens – 1 to 2 cm (0.4 to 0.8 in) diameter –[12] are roughly spherical and begin their development submerged in the ground, but gradually push above ground during maturation. In this state the outer surface is covered with mycelia, which forms a soft, fluffy coat that holds soil and debris to the outer surface.[8] The young fruit bodies often have a rounded knob or protuberance.[13] Like other members of genus Geastrum, G. pectinatum has a fruit body wall that is multilayered. At maturity, the outer layer (the exoperidium) splits open from the top in a stellate (star-shaped) manner into 7–9 rays that support the spore sac contained within the inner wall (the endoperidium). The expanded specimens are up to 5 cm (2.0 in) broad and 6 cm (2.4 in) tall.[10] The rays of the exoperidium bend back (reflex), simultaneously elevating the spore sac above the ground in what is known as the fornicate condition;[14] this position exposes the spore sac to more air currents, aiding spore dispersal.[15] The surface of the rays often crack to reveal lighter-colored areas, especially along the edges. Together with a well-developed layer of mycelium, the rays are typically bound to fragments of earth or forest duff.[10]

Close-up of pedicel and underside of spore sac

The tough and membranous endoperidium comprising the spore sac, purple-brown in color and 0.5 to 1.5 cm (0.2 to 0.6 in) tall by 1 to 2.5 cm (0.4 to 1.0 in) wide, is supported by a small stalk—a pedicel—that is 3–4 mm long by 7–10 mm wide and which has a grooved (sulcate) apophysis, or swelling. This ring-shaped swelling is made of remnants from a tissue called the pseudoparenchymatous layer.[16] When fresh, the pseudoparenchymatous layer is whitish in color, thick and fleshy; it dries to become brown to dark brown while shrinking and often splitting and peeling.[13] The endoperidium may be pruinose—covered with fine, white, powder—although the presence of this characteristic has been noted as being somewhat variable.[14][17] The spore sac is opened by a single apical pore atop a conical "beak", or peristome. The peristome is pectinate—made of tissue that resembles the teeth of a comb; the specific epithet is named after this characteristic. The peristome is 2 to 5 mm (0.08 to 0.20 in) long, and comprises 20–32 distinct ridges.[13] The mass of spores and surrounding cells within the sac, the gleba, is dark-brown, and becomes powdery in mature specimens. Internally, the endoperidium contains a structure called the columella that is narrowly conical in shape, whitish or pale brown, and extends more than halfway into the gleba.[13] G. pectinatum has no distinguishable odor or taste;[18] like other earthstar mushrooms, it is inedible,[19] and of "no alimentary interest".[20]

Microscopic characteristics

The spores of G. pectinatum are brown and opaque.[14] They have a roughly spherical shape[21] and are ornamented with transparent (hyaline), truncate warts;[12] the diameter is 4–4.5 µm, or 5.5–6.5 if the lengths of the warts is included.[22] Spore-bearing cells, the basidia, are 2- or 4-spored, while cystidia (specialized sterile cells that occur at the hymenial surface in some mushrooms) are absent.[18] The capillitia—a mass of thread-like sterile fibers dispersed among the spores—are light brown and 3–7 µm in diameter. They are tapered, thick-walled with a narrow interior, and either smooth or slightly encrusted.[13]

Similar species
Herbarium specimen

Geastrum pectinatum has been mistaken for the morphologically similar but smaller species G. schmidelii. The latter species lacks vertical striations on the basal portions of the endoperidium, and does not have a pseudoparenchymatous collar around the stem.[22] Another similar species, G. berkeleyi, has a shorter stem and is missing the ridges at the base of the spore sac.[23] Further, the color of its spore sac is usually brown, in contrast to the gray-blue of G. pectinatum.[24] G. xerophilum also has a dusting of white powder on the surface of the spore sac, but unlike G. pectinatum, consistently lacks a ring at the base of the pedicel; furthermore, in contrast to G. pectinatum, the spores of G. xerophilum are yellow and contain oil drops that are readily observable with a microscope.[14] G. striatum has smaller fruit bodies than G. pectinatum, and a distinct collar-like apophysis.[25]

Distribution and habitat

This species has been reported to grow solitary or in groups on sandy soil[21] or rich composted soil[26] in both mixed and coniferous forests, often beneath cedars.[27] In Hawaii, it is usually found growing in duff under coastal Casuarina and groves of Cupressus.[24] The species has been noted to occur in late summer and autumn (in Britain and Europe),[18] but the fruit bodies may dry and persist for some time.[10]

Geastrum pectinatum has a cosmopolitan distribution. It has been reported from Australia,[28] and New Zealand,[29] Africa (the Congo,[16] South Africa[16]) Central America (Costa Rica[30]), Asia (Northeastern China[31] and Japan[2]), and South America (Brazil[21]). In Europe, it has been reported from Belgium,[32] Ireland,[33] Germany,[34] the Netherlands,[35] Norway,[36] and Sweden.[37][38][39] In the Middle East, it has been recorded in Israel,[40] and Turkey.[41] In North America, it is known from the United States[12] (including Hawaii[14]), Canada,[10] and Mexico.[42] It is in the Red Data Book (documenting rare and endangered species) of Latvia,[43] and is considered a threatened species in Poland.[44] North American sources gives its frequency of appearance as "rare",[10][12] but Stellan Sunhede, in his 1989 monograph on the Geastraceae, considers it one of the most common earthstar mushrooms of northern Europe.[45]

Calcium oxalate crystals

Calcium oxalate is a common crystalline compound found in many fungi,[46] including the earthstars.[47] The presence of calcium oxalate crystals—apparent as a whitish powder on the surface of the spore sac—has been verified for G. pectinatum using scanning electron microscopy. The calcium oxalate crystals occur in the tetragonal form, known as weddellite.[48] A study on the related species Geastrum saccatum has shown that these crystals are responsible for the characteristic opening (dehiscence) of the outer peridial layers.[49] The formation of calcium oxalate crystals stretches the layers of the outer walls, pushing apart the inner and outer layers of the peridium.[49]

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References
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  2. Imai S. (1936). "Symbolae ad floram mycologicam Asiae Orientalis. I" [Contributions to the mycological flora of Eastern Asia]. Botanical Magazine (Tokyo). 50 (592): 216–24.
  3. Persoon CH. (1801). Synopsis Methodica Fungorum (in Latin). 1. Göttingen, Sweden: Apud H. Dieterich. pp. 132–3.
  4. Berkeley MJ, Curtis MA (1860). "Characters of new fungi, collected in the North Pacific Exploring Expedition by Charles Wright". Proceedings of the American Academy of Arts and Sciences. 4: 124.
  5. Berkeley MJ. (1839). "Descriptions of exotic fungi in the collection of Sir W.J. Hooker, from memoirs and notes of J.F. Klotzsch, with additions and corrections". Annals and Magazine of Natural History. 1. 3: 399. doi:10.1080/03745483909443251.
  6. Berkeley MJ. (1848). "Decades of fungi. Decade XX. Tasmanian fungi". London Journal of Botany. 7: 572–80.
  7. Palmer JT. (1959). "Observations on gasteromycetes. VIII. Persoon's specimens of Geastrum pectinatum Pers. and a reassessment of Geastrum plicatum Berk. and Geastrum tenuipes Berk". Persoonia. 1: 149–64.
  8. Ponce de Leon P. (1968). "A revision of the family Geastraceae". Fieldiana. Chicago, Illinois: Chicago Natural History Museum. 31 (14): 309, 327–8.
  9. Published in: Pilát A. (1958). Gasteromycetes, Houby-Břichatky. Flora ČSR B1 [Gasteromycetes, Puffballs] (in Czech). Prague, Czechoslovakia: Nakladatelstvi Československé Akademie Vĕd.
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  11. Gray SF. (1821). A Natural Arrangement of British Plants. London, UK: Cradock, and Joy. p. 585.
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  13. Laessoe T, Pegler DN, Spooner B (1995). British Puffballs, Earthstars and Stinkhorns: an Account of the British Gasteroid Fungi. Kew, UK: Royal Botanic Gardens. p. 90. ISBN 0-947643-81-8.
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  25. Sunhede (1898), p. 426.
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  34. Winterhoff W. (1981). "Old and new earthstar findings in the drift sand area between Walldorf and Mainz West Germany". Hessische Floristische Briefe (in German). 30 (2): 18–27.
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  38. Kers LE. (1977). "Some Gasteromycetes from Medelpad Sweden". Svensk Botanisk Tidskrift (in Swedish). 71 (1): 79–83.
  39. Andersson U-B. (2009). "Jordstjärnor i Sverige 1. Kamjordstjärna och kantjordstjärna" [Swedish earthstars (Gastraceae) I]. Svensk Botanisk Tidskrift (in Swedish). 103 (2): 113–5.
  40. Binyamini N. (1994). "New records of higher fungi from Israel". Mycoscience. 35 (4): 425–8. doi:10.1007/BF02268518.
  41. Kaya A. (2006). "Macrofungi from Andırın (Kahramanmaras) district". Turkish Journal of Botany. 30 (2): 85–93. ISSN 1300-008X.
  42. Vasquez LS, Guzmán-Dávalos L (1990). "New records of fungi macromycetes for the states of Jalisco Puebla and Zacatecas Mexico". Brenesia (in Spanish). 33: 61–74.
  43. Vimba E. (1997). "Mycological studies of the Latvian coast of the Baltic Sea and the Gulf of Riga". Proceedings of the Latvian Academy of Sciences Section B Natural Exact and Applied Sciences. 51 (5–6): 234–40.
  44. Wojewoda W. (2000). "New localities of rare and threatened species of Geastrum (Lycoperdales) in Poland". Acta Mycologia. 35 (2): 145–51.
  45. Sunhede (1898), p. 294.
  46. Whitney KD, Arnott HJ (1986). "Morphology and development of calcium oxalate deposits in Gilbertella persicaria (Mucorales)". Mycologia. 78 (1): 42–51. doi:10.2307/3793375. JSTOR 3793375.
  47. Horner HT, Tiffany LH, Cody AM (1983). "Calcium oxalate bipyramidal crystals on the basidiocarps of Geastrum minus (Lycoperdales)". Proceedings of the Iowa Academy of Sciences. 92 (2): 70–7.
  48. Krisai I, Mrazek E (1986). "Calcium oxalate crystals in Geastrum". Plant Systematics and Evolution. 154 (3–4): 325–41. doi:10.1007/BF00990131.
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Cited literature
  • 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.
  • botany.cz Geastrum pectinatum Pers. – hvězdovka dlouhokrká / hviezdovka dlhokŕčková (in Czech)

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