Gomphus clavatus

Gomphus clavatus
Scientific classification
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Phallales
Family: Gomphaceae
Genus: Gomphus
Species: G. clavatus
Binomial name
Gomphus clavatus
(Pers.) Gray (1821)[1]
Synonyms[2]
Gomphus clavatus
View the Mycomorphbox template that generates the following list

Mycological characteristics

ridges on hymenium
cap is infundibuliform
hymenium is decurrent
stipe is bare
spore print is yellow
ecology is mycorrhizal
edibility: edible

Gomphus clavatus, commonly known pig's ears or violet chanterelle, as is an edible species of fungus in the genus Gomphus, family Gomphaceae native to Eurasia and North America. The fruit body is vase- or fan-shaped with wavy edges to its rim, and grows up to 15 cm (6 in) wide and 17 cm (6 3⁄4 in) tall. The upper surface or cap is orangish-brown to lilac, while its lower spore-bearing surface, the hymenium, is covered in wrinkles and ridges rather than gills or pores, and is a distinctive purple color. Described by Jacob Christian Schäffer in 1774, G. clavatus has had several name changes and many alternate scientific names, having been classified as a chanterelle by several authorities, though it is not closely related to them.

Typically found in coniferous forests, G. clavatus is mycorrhizal, and is associated with tree species in a variety of coniferous genera, particularly spruces and firs. It is more common at elevations of greater than 2,000 ft (600 m), in moist, shady areas with plenty of leaf litter. Although widespread, G. clavatus has become rare in many parts of Europe and extinct in Great Britain. It has been placed on the national Red Lists of threatened fungi in 17 European countries and is one of 33 species proposed for international conservation under the Bern Convention.

Taxonomy

German naturalist Jacob Christian Schäffer described Elvela (subsequently Helvella) purpurascens in 1774.[3] Austrian naturalist Franz Xaver von Wulfen gave it the name Clavaria elveloides in 1781, reporting that it appeared in the fir tree forests around Klagenfurt in August and was common around Hüttenberg. He recorded that poor people ate it, giving it the local name hare's ear.[4] In 1796, mycologist Christian Hendrik Persoon described G. clavatus as Merulius clavatus, noting that it grew in grassy locations in woods. He noted it was the same species that Schäffer had described.[5] The specific epithet—derived from the Latin word clava (club) and meaning "club-shaped"[6]—refers to the shape of young fruit bodies.[7] In his 1801 Synopsis methodica fungorum, Persoon placed Merulius clavatus (recognising two varieties—violaceus and spadiceus) in a section Gomphus within Merulius.[8]

British botanist Samuel Frederick Gray used Persoon's name, transferring the violet chanterelle to the genus Gomphus in 1821.[9] As it is the first named member of the genus it became the type species.[9] The starting date of fungal taxonomy had been set as January 1, 1821, to coincide with the date of the works of the 'father of mycology', Swedish naturalist Elias Magnus Fries, which meant the name required sanction by Fries (indicated in the name by a colon) to be considered valid. Thus the species was written as Gomphus clavatus (Pers.: Fr.) Gray. A 1987 revision of the International Code of Botanical Nomenclature set the starting date at May 1, 1753, the date of publication of Linnaeus' work, the Species Plantarum.[10] Hence, the name no longer requires the ratification of Fries' authority.[2] Persoon followed suit in treating Gomphus as a separate genus in his 1825 work Mycologia Europaea.[9] Here he recognised M. clavatus as the same species as Clavaria truncata described by Casimir Christoph Schmidel in 1796, calling the taxon Gomphus truncatus.[11]

Fries himself declined to keep the genus separate,[9] instead classifying Gomphus as a tribe within the genus Cantharellus in his 1821 work Systema Mycologicum, the species becoming Cantharellus clavatus. He recognised four varieties: violaceo-spadiceus, carneus, purpurascens and umbrinus.[12] Swiss mycologist Louis Secretan described three taxa—Merulius clavatus carneus, M. clavatus violaceus and M. clavatus purpurascens—in his 1833 work Mycographie Suisse.[13] Many of his names have been rejected for nomenclatural purposes because Secretan had a narrow species concept, dividing many taxa into multiple species that were not supported by other authorities, and his works did not use binomial nomenclature consistently.[14][15] Fries revised his classification in his 1838 book Epicrisis Systematis Mycologici seu Synopsis Hymenomycetum, placing it in a series—Deformes—in the genus Craterellus.[16]

1897 illustration by Albin Schmalfuss

Paul Kummer raised many of Fries' tribi (subgenera) to genus rank in his 1871 work Der Führer in die Pilzkunde, classifying the violet chanterelle in the genus Thelephora.[17] Jacques Emile Doassans and Narcisse Théophile Patouillard placed it in the genus Neurophyllum (also spelt Nevrophyllum) in 1886,[18][9] removing it from Cantharellus on account of its orange spores. However, Charles Horton Peck discarded the name in 1887 and returned G. clavatus to Cantharellus.[19] In 1891, German botanist Otto Kuntze published Revisio generum plantarum, his response to what he perceived as poor method in existing nomenclatural practice.[20] He coined the genus Trombetta to incorporate the violet chanterelle, hence giving it the name Trombetta clavata.[21] However, Kuntze's revisionary program was not accepted by the majority of botanists.[20]

Alexander H. Smith treated Gomphus as a section within Cantharellus in his 1947 review of chanterelles in western North America, as he felt there were no consistent characteristics that distinguished the two genera.[22] In 1966 E.J.H. Corner described a small-spored variety, G. clavatus var. parvispora, from specimens collected in Uganda;[23] it is not considered to have independent taxonomic significance.[24]

Research combining the use of phylogenetic analyses of DNA sequences and more traditional morphology-based characters has resulted in a reshuffling of the species concept in Gomphus;[19] as a result, G. clavatus is considered the only Gomphus species in North America.[25] Comparison of the DNA sequences of species Gomphus brevipes and Gomphus truncatus has shown them to be genetically identical to G. clavatus, and they may be treated as synonyms.[19]

Gomphus clavatus is commonly known as pig's ears, alluding to the violet underside and yellowish cap of the fruit bodies,[26] although this vernacular name is also used for Discina perlata.[27] Other English common names for this species include clustered chanterelle and violet chanterelle.[28] Gray coined the name clubbed gomphe.[1] In the Sherpa language of Nepal the fungus is known as Eeshyamo ("mother-in-law"), as its imposing fruit body is reminiscent of a mother-in-law, who has a dominant role in the Sherpa family.[29]

Description

The wrinkled and ridged surface of the hymenium

The basidiocarps, or fruit bodies, of immature Gomphus clavatus are club-shaped and have one cap or pileus, but later spread out and have a so-called merismatoid appearance—several vase-shaped caps rising from a common stem. The fruit body is up to 15 cm (6 in) wide and 17 cm (6 3⁄4 in) tall, fan-shaped with wavy edges. The upper surfaces of the fruit bodies are covered with brown hyphae that form small, distinct patches towards the margin, but combine to form a continuous felt-like tomentum over the center of the cap. The color of the upper cap surface is orange-brown to violet, but later in age fades to a lighter brown.[30] The cap margins of older mushrooms can be quite ragged.[31] The lower spore-bearing surface—the hymenium—is wrinkled, often with folds and pits, and violet to brown in color. The solid stem, which is continuous with the cap,[32] is 0.8–3 cm (3⁄8–1 1⁄8 in) wide, 4–10 cm (1 5⁄8–3 7⁄8 in) tall,[22] and covered with fine hairs (tomentum) that become more coarse (hispid) towards the base. It is often compound, with several fruit bodies arising from the basal portion. Fruit bodies may bruise reddish-brown where handled.[33] The flesh can be whitish-pink to lilac or cinnamon-buff. Thick under the centre of the cap, it thins out towards the margins.[22] It can be crunchy, though it is softer than that of the chanterelle.[31]The taste and odor are mild. The spore print is yellow to orange-yellow.[34]

Basidiospores are elliptical, wrinkled or slightly warted, and 10–14 by 5–7.5 μm.[30] The spores are nonamyloid, meaning they have a negative color reaction with the iodine in Melzer's reagent. The spore-bearing structures, the basidia, are elongated or club-shaped, hyaline (glassy or translucent), and four-spored, with dimensions of 60–90 by 8.5–11.5 μm.[35] G. clavatus does not contain cystidia, the sterile cells associated with basidia in many species. Clamp connections are present.[35]

Similar species

Gomphus crassipes, found in Spain and North America, can only be reliably distinguished from G. clavatus with the use of a microscope.[36] Its basidiospores are generally longer (11-17 by 5.5-7 μm) and have a more finely wrinkled surface.[19] Pseudocraterellus pseudoclavatus (formerly classified in Gomphus) is a lookalike species that grow under conifers in the central United States and westward,[37] also differing on microscopic characters and reaction to potassium hydroxide.[22] Turbinellus floccosus and T. kauffmanii are of similar shape but their caps are covered in scales.[7]

Habitat, distribution, and conservation

Gomphus clavatus fruiting in a fairy ring

Gomphus clavatus grows singly, in clusters or clumps, or even occasionally fairy rings,[22] on the ground, typically in coniferous forests, and with a preference for moist, shady areas with deep leaf litter,[34] or rotten wood debris on the ground. The species is equally common in older or younger stands of trees.[19] Fruit bodies are easily missed because their colors blend with those of the forest floor.[38] It is more common at elevations of greater than 2,000 ft (600 m).[28] Gomphus clavatus has been reported as forming symbiotic (mycorrhizal) associations with a variety of trees: Abies alba,[39] Abies cephalonica,[19] Abies firma,[40][41] Abies nephrolepis,[42] Abies religiosa,[43] Picea species,[44] Pinus densiflora,[40][41] Pseudotsuga menziesii,[45] and Tsuga heterophylla.[45][46] It is also reported with beech (Fagus sylvatica) in Europe.[47]

In Asia, Gomphus clavatus has been reported from China,[23] Japan,[23] Korea,[48] Malaysia,[49] Nepal,[29] and Pakistan.[23] European countries that have reported the fungus include Austria,[50] the Czech Republic,[51] France,[18] Germany,[52] Greece,[50] Italy,[50] Lithuania,[53] Poland,[54] Romania,[55] Russia,[45] Sweden,[26] Switzerland,[50] and Turkey.[56] In North America, the fungus has been found across Canada,[23] Mexico,[50] and the United States,[35] where it is abundant in the Pacific Northwest.[22]

In Europe, Gomphus clavatus appears on the national Red Lists of threatened fungi in 17 countries and is one of 33 species proposed for international conservation under the Bern Convention.[47] Due to a substantial decline in sightings,[57] Gomphus clavatus became a legally protected species in Hungary on September 1, 2005.[58] It also has legal protection in Slovakia[47] and Slovenia.[59] The species formerly occurred in England, but has not been seen since 1927 and is now regarded as extinct.[60] The fungus faces loss and degradation of its habitat; eutrophication is another potential threat.[47] Gomphus clavatus was selected as the 1998 Pilz des Jahres ("Mushroom of the Year") by the German Mycological Society, partly to highlight its vulnerable status.[61]

Edibility

Gomphus clavatus is edible (and rated as choice by some),[34][62] while others find it tasteless.[63] It has an earthy flavor and meaty texture that complements red meat dishes.[38] Like many edible fungi, consumption may cause gastrointestinal distress in susceptible individuals.[33] The flesh becomes bitter with age,[34] and older specimens may be infested with insects.[63] Insect infestation is unlikely if the weather is cool.[64] G. clavatus has been used for cooking for some time—Fries included it in his 1867 book Sveriges ätliga och giftiga svampar (Edible and Poisonous Mushrooms in Sweden). It is highly regarded by the Zapotec people of Ixtlán de Juárez in Oaxaca,[65] and the Sherpa people in the vicinity of Sagarmatha National Park in Nepal.[29]

Extracts prepared from G. clavatus fruit bodies have a high antioxidant activity,[66] and a high concentration of phenolic and flavonoid compounds.[67] Phenolic compounds identified from the fungus include protocatechuic acid, gallic acid, gentisic acid, vanillic acid, syringic acid, cinnamic acid, caffeic acid, ferulic acid, and tannic acid.[68] In a chemical analysis of collections from the South Aegean Region of Turkey, the fungus was shown to have bioaccumulated the toxic metal cadmium to levels exceeding the maximum intake recommended by the European Union Scientific Committee on Food.[67]

References

  1. 1 2 Gray SF. (1821). A Natural Arrangement of British Plants 1. London, UK. p. 638.
  2. 1 2 "Gomphus clavatus (Pers.) Gray". MycoBank. International Mycological Association. Retrieved 17 December 2015.
  3. ↑ Schäffer JC. (1774). Fungorum qui in Bavaria et Palatinatu circa Ratisbonam nascuntur Icones (in Latin) 4. Erlangen, Germany: J.J. Palmium. p. 109.
  4. ↑ Jacquin NJ. (1781). Miscellanea Austriaca ad Botanicum, Chemiam et Historiam Naturalem Spectantia (in Latin) 2. Vienna: J.P. Kraus. p. 99; plate 12:1.
  5. ↑ Persoon CH. (1795). "Observationes Mycologicae". Annalen der Botanik (Usteri) (in Latin) 15: 1–39 (see p. 21).
  6. ↑ Stearn WT. (2004). Botanical Latin. Oregon: Timber Press. p. 386. ISBN 978-0-88192-627-9.
  7. 1 2 Roody WC. (2003). Mushrooms of West Virginia and the Central Appalachians. Lexington, Kentucky: University Press of Kentucky. p. 135. ISBN 978-0-8131-9039-6.
  8. ↑ Persoon CH. (1801). Synopsis Methodica Fungorum [Methodical Synopsis of the Fungi] (in Latin) 2. Göttingen, Germany: Apud H. Dieterich. p. 498.
  9. 1 2 3 4 5 Giachini AJ, Castellano MA. (2011). "A new taxonomic classification for species in Gomphus sensu lato". Mycotaxon 115: 183–201. doi:10.5248/115.183.
  10. ↑ Esser K, Lemke PA. (1994). The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research. Heidelberg, Germany: Springer. p. 81. ISBN 978-3-540-66493-2.
  11. ↑ Persoon CH. (1825). Mycologia Europaea (in Latin) 2. Erlangen, Germany: Palm. p. 9.
  12. ↑ Fries EM. (1821). Systema Mycologicum (in Latin) 1. Lund, Sweden. p. 322.
  13. ↑ Secretan L. (1833). Bonnant PA., ed. Mycographie suisse, ou, Description des champignons qui croissent en Suisse, particulièrement dans le canton de Vaud, aux environs de Lausanne [Swiss Mycographie, or description of fungi growing in Switzerland, particularly in the canton of Vaud, near Lausanne] (in French) 2. Geneva, Switzerland. pp. 471–73.
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  18. 1 2 Doassans MME, Patouillard NT. (1886). "Champignons du Béarn (2eliste)". Revue Mycologique (in French) 8: 25–28. (Reprint from "Collected Mycological Papers" chronologically arranged and edited by L. Vogelenzang, Librarian Rijksherarium, Leiden, Amsterdam, vols. 1–3, 1978)
  19. 1 2 3 4 5 6 Giachini A. (2004). Systematics, Phylogeny, and Ecology of Gomphus sensu lato (Ph.D. thesis). Corvallis, Oregon: Oregon State University.
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  47. 1 2 3 4 Dahlberg A, Croneborg H. (2006). The 33 Threatened Fungi in Europe (Nature and Environment). Strasbourg, Germany: Council of Europe. pp. 58–60. ISBN 978-9287159281.
  48. ↑ Han S-K, Park Y-J, Choi S-K, Lee J-O, Choi J-H, Sung J-M. (2006). "Some unrecorded higher fungi of the Seoraksan and Odaesan National Parks". Mycobiology 34 (2): 56–60. doi:10.4489/MYCO.2006.34.2.056.
  49. ↑ Jais HM, Tajuddin R, Iffendy KA. Macrofungi of a Healthy Campus. Penerbit USM. p. 41. ISBN 978-983-861-869-4.
  50. 1 2 3 4 5 Petersen RH. (1971). "The genera Gomphus and Gloeocantharellus in North America". Nova Hedwigia 21: 1–118.
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  52. ↑ Dörfelt H, Bresinsky A. (2003). "Distribution and ecology of selected macromycetes in Germany" (PDF). Zeitschrift für Mykologie 69 (2): 177–286 (see p. 200).
  53. ↑ Urbonas VA, Matyalis AA, Gritsyus AI. (1990). "Trends of variability of macromycetes, extinguishing species and principles of their protection in Lithuania". Mycology and Phytopathology 24: 385–88.
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  55. ↑ Pop A, Soltesz AM. (2001). "Mushrooms from the Barsa Depression (Brasov County)". Contributii Botanice 36: 41–51.
  56. ↑ Sesli E. (1997). "Two new records of cantharelloid fungi for Turkey". Israel Journal of Plant Sciences 45: 71–74. doi:10.1080/07929978.1997.10676672.
  57. ↑ Siller I, Vasas G, Pal-Fam F, Bratek Z, Zagya I, Fodor L. (2005). "Hungarian distribution of the legally protected macrofungi species". Studia Botanica Hungarica 36: 131–63.
  58. ↑ Siller I, Dima B, Albert L, Vasas G, Fodor L, Pal-Fam F, Bratek Z, Zagya I. (2006). "Protected macrofungi in Hungary". Mikologiai Kozlemenyek 45: 3–158.
  59. ↑ Al-Sayegh Petkovsek S, Pokorny B, Piltaver A. (2004). "The first list of macrofungi from the wider area of the Salek Valley". Zbornik Gozdarstva in Lesarstva 72: 83–120.
  60. ↑ "Gomphus clavatus (Pers.) Gray, Nat. Arr. Brit. Pl. (London) 1: 638 (1821)". Checklist of the British & Irish Basidiomycota. Royal Botanical Gardens, Kew. Retrieved 15 September 2011.
  61. ↑ "1998: Gomphus clavatus (Pers.: Fr.) S.F.Gray, Schweinsohr" (in German). Deutsche Gesellschaft für Mykologie. Retrieved 15 December 2015.
  62. ↑ Weber NS, Smith AH. (1980). The Mushroom Hunter's Field Guide. Ann Arbor, Michigan: University of Michigan Press. p. 84. ISBN 978-0-472-85610-7.
  63. 1 2 Davis RM, Sommer R, Menge JA. (2012). Field Guide to Mushrooms of Western North America. Berkeley, California: University of California Press. p. 275. ISBN 978-0-520-95360-4.
  64. ↑ Smith AH, Weber NS. (1980) [1958]. The Mushroom Hunter's Field Guide. University of Michigan Press. p. 83. ISBN 978-0-472-85610-7.
  65. ↑ Garibay-Orijel R, Caballero J, Estrada-Torres A, Cifuentes J. (2007). "Understanding cultural significance, the edible mushrooms case". Journal of Ethnobiology and Ethnomedicine 3 (4): 4. doi:10.1186/1746-4269-3-4.
  66. ↑ Makropoulou M, Aligiannis N, Gonou-Zagou Z, Pratsinis H, Skaltsounis A-L, Fokialakis N. (2012). "Antioxidant and cytotoxic activity of the wild edible mushroom Gomphus clavatus". Journal of Medicinal Food 15 (2): 216–21. doi:10.1089/jmf.2011.0107.
  67. 1 2 Sarikurkcu C, Tepe B, Kocak MS, Uren MC. (2015). "Metal concentration and antioxidant activity of edible mushrooms from Turkey". Food Chemistry 175: 549–55. doi:10.1016/j.foodchem.2014.12.019.
  68. ↑ Puttaraju NG, Venkateshaiah SU, Dharmesh SM, Urs SMN, Somasundaram R. (2006). "Antioxidant activity of indigenous edible mushrooms". Agricultural and Food Chemistry 54 (26): 9764–72. doi:10.1021/jf0615707.

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