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Species Diversity of the Genus Psilocybe (Basidio- Mycotina, Agaricales, Strophariaceae) in the World Mycobiota, with Special At

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Species Diversity of the Genus Psilocybe (Basidio- Mycotina, Agaricales, Strophariaceae) in the World Mycobiota, with Special At International]ournal ofi11edicinal J11ushrooms, Vol. 7, pp. 305-331 (2005) Species Diversity of the Genus Psilocybe (Basidio­ mycotina, Agaricales, Strophariaceae) in the World Mycobiota, with Special Attention to Hallucinogenic Properties Gastón Guzmán In~tituto de Ecología, Apartado Postal 63, Xalapa 91000, Veracruz, Mexico; [email protected] This article is dedícated to the outstanding mycologist, colleague, andfriend, Professor Shu-Ting Chang ABSTRACT: An exhaustive world revision of ali names considered in the genus Psilocybe s.l. is pre­ sented, of which the haliucinogenic species were treated with special emphasis. Seven hundred eighteen names related to Psilocybe were found reportcd in the bibliography, of which only 227 are accepted taxa in P;ilqcybe. The concept of the genu~ foUowed here is that of Guzmán 1983; th ~i:~fore Hypholo"!:ª' lVlelanótus, and Stropharza were excluded: Moreover, 53 spec1es of Psathyrella, m8.ny times related w1th Psilocybe, were also e..'Ccluded. The hallucinogenic species are 144, which are oistübuted m ali the coriti­ nents, of which Latín America (iiiduding the Caribbae), has the top, with more than 50 species. There are only 22 species in Canada and the US, while Mexico is the country with the higest number iri t he world, with 53 species. Europe has only 16 species, Asia 15, Africa 4, and Australia·and eastern islands 19. Sorne Psilocybe·species are common iii several countries or regions, as are P cubensis and P subcuben­ sis in ali the tropics; P coprophila in many temperate and tropical regions; P. argentina in several high ___ _111ountains or in tl:ie Austral anci .B_on;al_ regiQn§; anq fjin1etcy:ja and P semi/aneeata in Et!JOp_e, Carni,da, ----~~ -~--- e - .. and the US, but U:nknown in Mexi~o. ... - .... KEYWORDS: Psilocybe, species diversity, hallucinogenic species, world distribution INTRODUCTION des and P semilanceata are only recognized, but P montana was confusedly related with P atrorufa; Since the fi.rst descriptions and concepts of Psi­ P merdaria was considered in Psalliota (Fr.) P. locybe (Fr.) P. Kumm. (e.g., Fries, 1821, 1836; Kumm. (a synonym of Agaricus L.), and Psilocybe Kummer, 1871; Qi.élet, 1888; Ricken, 1915), it luteonitens and P squamosa were coO:sidered also in was involved with several genera, such as Agaricus Psalliota. Kummer (1871) and Ricken (1915) rec­ L., Agrocybe Fayod, Hypholoma (Fr.) P. Kumm., ognized the genus Psilocybe, but still they confused Panaeolina Maire, Panaeolus (Fr.) Qi.él., Psathy- several genera in it. Qy.elét (1888) described the 1·ella (Fr.) ~él. and Stropharia (Fr.) ~él., among genus Geophila Qy.él., where he involved Psilocybe, others. Fries (1821) described 22 taxa as Agaricus, Psathyrella and others. Kühner and Romagnesi tribus Psilocybe Fr., of which only two are now (1953) followed ~elét. recognized in Psilocybe: P merdaria and P montana. 1he modern studies in the genus began mainly Later, Fries (1836) described 27 species, of which with Singer (1949), but the interest in the genus P atrobrunnea, P bu/lacea, P coprophila, P physaloi- started with the descovery of the hallucinogenic 1521-9437/05 $35.00 © 2005 by Begell House, Inc. 305 i ! 1 ¡-· G.GUZMÁN j ' ;. _J. fungi in Mexico, through the works ofWasson and Agaricales, divided the genus Psilocybe in Psilocybe ; Wasson (e.g., Wasson and Wasson, 1957). s.s. and psychedelia, to be considered in this last Heim (19 56) published the fust taxonomic record group the hallucinogenic species of Psilocybe. The of the hallucinogenic Psilocybe in Me.xico gathered by author reserves his opinion on this new genus in the vVassons, although Schultes (1939) and Singer another paper. (1949, pp. 472 and 506) briefiy and confusedly had Previous to this report, Guz~~n and Vergeer reported these mushrooms. H owever, the taxonomic (1978) presented a checklist of 390 names consid­ studies in Psilocybe were consolidated with the works ered in Psilocybe, with their bibliographic reference ofHeim and Wasson (1958) and Singer and Smith in each one. The present paper is part of the second (1958), both on the hallucinogenic species. edition of1he Genus Psilocybe, which the author has Guzmán (1983) wrote the fustworld monograph in preFaration to publis,h through Koeltz Scientific on the genus, with 144 accepted taxa of 467 names Books (Germany). reported in the bibliography. However, he described 52 taxa as new. The hallucinogenic species considered by Guzmán were 91 taxa (no. 81 was reported by THE TAXA O F PSILOCYBE .R EP ORTEO IN mistake). Later, Guzmán (1995) published a supple­ THE BIBLIOGRAPHY ment with 29 _additional taxa described by other specialists including himself, and 5 new species, A careful revision of all the species qr names After that, Guzmán and his colleagues published reported in Psilocybe or related to it by the bibli- several papers with more newspecies,hallucinogenic ographywas made, based :first in Guzmán (1983, andnothallucinogenlcm~hrootµs(G~zmánetal., 1995), Guzmán ;and Vergeer (1978), and ~n ali· 1997a,b, 1999, Z002, 2003 ~)); 2004a,b~ Gu:~min, the available wO,rkS discussed above, inclÜ,ding · 1998,2000,2004a,2005; Guzmán ~nd Castro,2003; . Re:i. (1922), Smith (1908), and others," as well as Guzmán and Kasuya, 2004). in the Index Fungorum (Kirk, 2004), as shown in Also Johston and Buchanan (1995), Reid and Appendix 1 (see page 313). There are 718 names, Eicker (1999), and 1homas et al. (2002) described of which only 227 are recognized in Psilocybe s.s. new species. Singer (1986) considered 86 species Kirk (2004) reported 506 names, but 51 were _, in Psilocybe. Watling andGregory(:L9.3-.'4-.dis.~.?_ed . ~...:epr.ated b·ecause he. :considered twice. the type ali the species of Psilocybe in Great Britain. Stamets species in those cases where there are varieties or (1996) presented a compilation of the knowledge forms (example, Psilocybe agrariella one time and of the genus in the world. Noordeloos (1999, 2001) P. agrariella var. agrariella another). It is interesting revised the Neerlandica species and a taxonomic to observe that after 1983 there was an increase of study of the section Psilocybe s. Singer. N oordeloos 83 accepted species in the genus. in the above papers considered Psilocybe p.p: in the Guzmán (1995) considered Psilocybe divided Friesian sense to include Hypholoma, Melanotus in 16 sections, of which Cordiporae and Semila- Pat., and Stropharia, criterion not followed here. nceatae presented the highest number of species, Boekhout et al. (2002) made molecular studies with around 20 for each. However, the classifica- in the section Psilocybe s. Singer. Bon and Roux tion of Singer (1986) distinguished seven sections: (2003) revised in a key ali the European species of Nlerdariae (Fr.) Singer, Caerulescentes Singer, Tenaces Psilocybe s.s., and Krieglsteiner (1984, 1986) and (Fr.) Sacc., Atrobrunnea Singer, Septembres Singer, Gartz (1998) revised the complex P. cyanescens both Psilocybe, and Chrysocystidiatae Singer, ofwhich Cae- in Europe and North America. All the contribu- rulescentes and Psilocybe, with 34 and 18, respectively, tions of Pegler (e.g., Pegler, 1983, 1989, 1997) were are the bigest sections. considered in this paper. Sorne species are recognized in the present paper Other important revised works were Heim et al. in a concept different from that of Guzmán (1983, (1967), Smith (1949), and Wu et al. (1997). Recently, 1995), in order to follow sorne recent taxonomic or Moncalvo et al. (2002) in a molecular study of several nomenclatural changes as those of Singer (1986), • /nternationa/ Journal of Medicinal Mushrooms 306 SPECIES DIVERSITYIN GENUS PSILOCYBE vVatling and Gregory (1987), Noordeloos (1999, are 31, 4, and 13, respectively. There are 53 species . 2001), and Bon and Roux (2003).1hese species are of Psathyrella described as Psilocybe. Other genera P alfofimbriata, P caeruleoannulata, P callosa, P chi­ confusedly related to Psilocybe areAstylospora Fayod onophila, P fagicola, P strictipes, P thrausta, P velifera, (asAtylospora by Murrill), Clitopilus (Fr. ex Rabenth.) P xalapensis, and P wassoniorum. P. Kumm., Conocybe Fayod, Ca/erina Earle, Galeropsis Psilocybe coprophila, P cubensis, and P subcubensis Velen., Kuehneromyces Singei- et A.H. Smith, Lac­ are the most common species; they are reported rymaria Pat., JV1arasmiellus Murrill, Mythicomyces from a1l the continents. Certainly they are fimicolous Readhead et AH. Smith,Naucoria (Fr.) P. Kumm., fungi, but P cubensis and P subcubensis grow only in Phaeogalera Kühner, Phaeomarasmius Scherff., Pho­ tropical and subtropical regions, and P coprophila in !iota (Fr.) P. Kumm., Pholiotella Speg., and Tubariq. ali the regions. However, P merdaria, P jimetaria, P (W.G. Smith) Gillet. moelleri and P su!jimetaria, also funicolous species, have distribution restricted to temperate regions. Another fimicolous fungus is P argentina, which THE HALLUCINOGENIC SPECIES AND grows only in the Austral and Boreal regions of the THEIR DISTRIBUTION world or in the high mountains of the temperate, tropical, and subtropical regions. 1he k:nown hallucinogenic or neurotropic species 1here are still taxonomic problems in several in Psilocybe are 144 (Appendix 1), which represent species, mainly in P cynescens complex, divided in an increase of 53 species in comparison with those the European species: P cyanescens s.s., P arcana, reported in 1983. 1he criterion to distinguish th.e P bohemica, P liniformam var. liniformans, and P neurotrbpic species used by Guzrrúin (1983) con-­ serbica, arid in the North African species: P maire, sidered t.he bluing (caerulescent) to black:ish feature and in the complex from the US as P azurescens · in the basidioma, as well as the farinaceous taste and the American P cyanescens. It is probable that and odor. However, these features-the bluing a~d the P cyanescens reported from the US is a complex blackish-are not easy to find, because they depend of species. on the development of the basidioma. For instance, in P fagicola, P galindii, P mexicana, and P.
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