Acta Botánica Mexicana 100: 79-106 (2012) NEW TAXONOMICAL AND ETHNOMYCOLOGICAL OBSERVATIONS ON PSILOCYBE (FUNGI, BASIDIOMYCOTA, AGARICOMYCETIDAE, S.S. AGARICALES, STROPHARIACEAE) FROM MEXICO, AFRICA AND SPAIN Gastón Guzmán Instituto de Ecología, A.C. Apdo. postal 63, 91070 Xalapa, Ver., México [email protected] ABSTRACT A new emendation ofPsilocybe zapotecorum is presented, based on a new study on the holotype and on the description of pseudocystidia, pleurocystidia and cheilocystidia, An confusingly described previously. extensive and careful review of all holotypes of synonyms, and the wide neotropical distribution is also presented. The traditions as sacred mushroom are analyzed based on a study of códices and anthropological pieces, where P. zapotecorum and related species as P. muliercula and P. moseri are identified. Some ofthe códices and anthropological pieces showed that the cult of Quetzalcóatl was related to the use of these mushrooms. The application and meaning of the words “teonanácatl” from Sahagún and “teotlaquilnanácatl” by Guzmán are reviewed. Moreover, the relationship of P. mairei to the prehistoric mushroom depictions at Tassili in the Sahara Desert is discussed, and the first study of authentic material of this species from Algeria is presented, from where a neotype is selected. Finally, the relationship between a prehistoric mural in Spain and P. hispánica is also discussed. Key words: prehistoric muráis, Psilocybe hispánica, Psilocybe mairei, Psilocybe moseri, Psilocybe muliercula, Psilocybe zapotecorum, Quetzalcóatl, teotlacuilnanácatl. RESUMEN SepresentaunanuevaemendacióndePsilocybezapotecorum,basadaenunre-estudio del holotipo y en la descripción de los pseudocistidios, pleurocistidios y queilocistidos, 79 Acta Botánica Mexicana 100 79-106 2012 : ( ) antes confusamente descritos. Se discute la numerosa sinonimia de este hongo, con base en el estudio de los holotipos y su amplia distribución en el neotrópico. Se analizan sus usos tradicionales como hongos sagrados y se hace una revisión de códices y piezas antropológicas, en donde se identifica a esta especie o sus afines: P. muliercula y P. moseri. Se establece una relación entre el culto de Quetzalcóatl y el uso de tales hongos. Se analizan las palabras “teonanácatl” de Sahagún y “teotlaquilnanácatl” de Guzmán. Se discuten las relaciones de P. mairei con los murales prehistóricos de Tassili en el Desierto del Sahara. Se presenta el primer estudio de materiales auténticos de P. mairei de Argelia, de donde se designa un neotipo. Finalmente se tratan las relaciones entre un mural prehistórico en España y P. hispánica. Fa\abYasc\aYQ:nmra\QspYehistÓYÍcos,Psilocybehispanica,Psilocybemairei,Psilocybe moseri Psilocybe muliercula Psilocvbe zapotecorum, Quetzalcóatl, “teotlacuilnanácatl”. , , INTRODUCTION Despite the fact that the genus Psilocybe (Fr.) P. Kumm. has been known since the 18th century (Guzmán, 1983), it was only at the end ofthe 1950’s that an interest in knowing its species began, this in relation to the discovery of the hallucinogen- ic species in México. Many important studies have been published, e.g. Heim and Wasson (1958), Singer and Smith (1958), Heim et al. (1966), Guzmán (1978a,b, 1983, 1995, 2000, 2009), Wasson (1980), Redhead et al. (2007), and Noordeloos (2011), among others. However, despite extensive research on this genus, it is surprising that several taxonomic problems still remain, and the resolution of some ofthese is waiting, as well as the presentation ofnew ethnomycological data. This is the basis of the present article. In this way, a new concept ofPsilocybe zapotecorum is pro- vided, as well as its numerous synonyms, with a wide neotropical distribution, and its interesting relationships with Quetzalcóatl cult in ancient traditions of México. Also discussions on the identification ofthe mushrooms depicted in two prehistoric muráis in Africa and in Spain are considered. MATERIAL AND METHODS This paper is based on an intensive bibliographic review, related to the de- scriptions of Psilocybe zapotecorum under several ñames, by Heim, Singer and 80 Guzmán: New taxonomical and ethnomycological observations onPsilocvbe Smith, and Guzmán, among others, as well as on the ethnomycological information on this complex. Microscopic studies of several herbarium specimens, including the holotypes ofP. zapotecorum and related species were made. These microscopic observations were made with hand sections of the basidiome and mounted in 5% solution of KOH, with or without 1% Congo red solution on the slide, previously treated with 96% alcohol for rehydratation. In this way more than 300 exsiccata were checked from several herbaria, including the holotypes. The ñame neurotropic is used for these mushrooms, instead of the common ñame hallucinogenic, because it is more appropriate, as it was previously discussed by the author (Guzmán, 2009). RESULTS New concept ofPsilocybe zapotecorum its synonyms, distribution and traditions , in Mesoamerica and South America (Figs. 1, 4, 9-10, 14-24, 32) Following the new status of the genus Psilocybe (Fr.) R Kumm. proposed by Redhead et al. (2007), and accepted by the International Nomenclature Committee for Fungi (Norvell, 2007), Psilocybe s.l. is now divided in two genera: Psilocybe s.s. and Deconica (W.G. Sm.) P. Karst. In this way, all the bluing and then neurotropic species are included in Psilocybe s.s., whereas the non-bluing non-neurotropic are ac- commodated in Deconica. Concerning Psilocybe zapotecorum R. Heim (Figs. 1, 9), this species needs revisión, because it presents great variation in its macroscopic fea- tures in the color and form ofthe basidiome, as well as in the microscopic characters. After the first descriptions of P. zapotecorum from the Zapotee región in Oaxaca (México) by Heim (1956, 1957a), Heim and Wasson (1958), and Heim et al. (1966), as well as the publication of a color píate by Heim in Wasson (1957) and the Latin diagnosis in Heim (1957b), the identity of this mushroom was con- fused and several ñames were applied by some authors, as discussed below. First, Singer (1958) mistakenly redescribed this species based on specimens that Guzmán sent him at LIL as P. zapotecorum aff, gathered in 1958 from the type locality re- ported by Heim (1956). Guzmán’s material is Guzmán 1501-A at ENCB and LIL. The habitat ofthis collection, as that reported by Heim (e.g. Heim, 1956; Heim and Wasson, 1958), and discussed further by Guzmán (1978a) is swampy and muddy soils. Also, those mushrooms from the type locality were identified by local Zapo- tee people as “corona de Cristo” (Christ chrown), one of the common ñames of P. zapotecorum (Guzmán, 1997). Singer (1958) reported Guzmán’s fungus as a topo- 81 Acta Botánica Mexicana 100 79-106 2012 : ( ) type of P. zapotecorum. However, Guzmán’s collection is in fact P. hoogshagenii R. Heim (Fig. 7), a neurotropic and sacred mushroom described by Heim (Heim and Wasson, 1958) from Coatlán, in the Mixe zone, of Oaxaca, as stated Guzmán (1978a). Psilocybe zapotecorum and P. hoogshagenii are different species distin- guished by macro- and microscopio features (Guzmán, 1983). Independent ofthe preceding confusión, Singer and Smith (1958) described & P. candidipes Singer A.H. Sm. from Oaxaca as a neurotropic mushroom among & the Mazatecs. Also they described in the same article P. aggericola Singer A.H. Sm. from Argentina; moreover Singer determined P. aggericola var. alvara- doi Singer based on an herbarium specimen also from Argentina at BAFC. Later Heim (Heim et al., 1966) described P. zapotecorum f. elongata R. Heim from Oaxaca; Guzmán (1968) described P. bolivarii Guzmán from Sinaloa; Cifuentes and Guzmán (1981) described P. barrerae Cifuentes & Guzmán from Guerrero; and Guzmán (1982) described P. sanctorum Guzmán from the State of Méxi- co. From Brazil, Guzmán et al. (1984) described P. microcystidiata Guzmán & Bononi and P. zapotecorum var. ramulosum Guzmán & Bononi. Guzmán (1999) presented an emendation of P. barrerae, based on new observations of the pleu- rocystidia (now pseudocystidia), which were not considered in the original de- scription by Cifuentes and Guzmán (1981). Later, Guzmán (2000) described P. subzapotecorum Guzmán from Oaxaca, and more recently Guzmán et al. (2004) & described P. chaconii Guzmán, Escalona Ram.-Guill. from Veracruz. All ofthe above 12 ñames, are now merely synonyms ofP. zapotecorum as Guzmán (1983) previously noted for P. candidipes, P. aggericola var. aggericola, P. aggericola var. alvaradoi, P. zapotecorum f. elongata and P. bolivarii. The others are pro- posed as synonyms here for first time. These conclusions play an important role in the taxonomic concept of P. zapotecorum, and are the result of careful studies of the holotypes, where it was found that the pseudocystidia had not been previously considered, except for P. subzapotecorum. These pseudocystidia were mistakenly described as pleurocystidia in P. candidipes and P. aggericola by Singer and Smith (1958). Also, it was found that the form and color ofthe basidiome are highly variable and without taxonomic valué, as well as the form and size ofthe cheilocystidia. All these mushrooms were compared with others of the same section Zapotecorum Guzmán, e.g. P. moseri & Guzmán, P. muliercula Singer A.H. Sm. and others, in order to better define the limits ofP. zapotecorum, and the section Zapotecorum. Based on the confusión discussed above in the concept ofP. zapotecorum, a new emendation is presented. 82 Guzmán: New taxonomical and ethnomycological observations on Psilocybe Figs. 1-8. Important ethnomycological species ofPsilocybe in the world. 1. P. zapotecorum P (México); 2. mexicana (México); 3. P. caerulescens (México); 4. P. muliercula (México); 5. P. hispánica (Spain); 6. P. mairei (Africa); 7. P. hoogshagenii (México); 8. P. aztecorum (México). Scale bar: 1-2,4-7 = 20 mm, 3,8= 10 mm. 83 Acta Botánica Mexicana 100 79-106 2012 : ( ) Psilocybe zapotecorum R. Heim emend. nov. Holotype: Heim & Wasson No. J-125, August 3, 1956 (PC, isotypes ENCB, XAL) = P. zapotecorum R. Heim, Comp. Rend. Séan. PAcad. Se. 242: 1393, 1956, nom. nud. = P. zapotecorum R. Heim, Rev. Mycol. 22: 77, 1957. = P. zapotecorum emend. Guzmán, Nova Hedwigia 29: 633, 1978. = & P. candidipes Singer A.H. Sm., Mycologia 50: 141, 1958. = & P. aggericola Singer A.H. Sm., Mycologia 50: 142, 1958. = P. zapotecorum f. elongata R. Heim, Comp. Rend. Séan. PAcad. Se. 250: 1158, 1960, nom. nud. = P. aggericola var. alvaradoi Singer, in BAFC, 1965, nom. nud. = P. bolivarii Guzmán, Ciencia (Méx.) 26: 25, 1968. = P. barrerae Cifuentes & Guzmán, Bol. Soc. Mex. Micol. 16: 52, 1981! = P. sanctorum Guzmán, Bol. Soc. Mex. Mic. 17: 90, 1982! = P. microcystidiata Guzmán & Bononi, Mycotaxon 19: 345, 1984! = P. zapotecorum var. ramulosum Guzmán & Bononi, Mycotaxon 19: 346, 1984! = P. barrerae Cifuentes & Guzmán emend. Guzmán, Acta Bot. Mex. 49: 43, 1999. = P. subzapotecorum Guzmán, Doc. Mycol. 29 (116): 46, 2000! = P. pseudozapotecorum Guzmán, Doc. Mycol. 29 (116): 47, 2000, a typogra- phic mistake. = P. chaconii Guzmán, Escalona & Ram.-Guill., Int. J. Med. Mushrooms 6: 276, 2004! Thosefeatures in italic are new additions to the concept ofthe species. mm Pileus (20-)40-70(-110) diam., polymorphic, conic to convex, convex- plane or campanulate, regular or irregular, sometimes papillate, or subumbilicate, hygrophanous, yellowish palé to chocolate-brown, orangish-brown or cinnamon- brown, smooth, sublubricous, sometimes with white floccose scales from the veil at the margin. Lamellae adnexed to sinuate, whitish-brown or palé reddish-brown to dark violaceous, edges whitish. Stipe (70-)100-180(-200) x (5-)10-15(-20) mm, taper- ing upward, solid to hollow, fibrous, whitish to concolorous with pileus, covered by short or large, floccose, white appressed scales toward the base, frequently in multi- 84 Guzmán: New taxonomical and ethnomycological observations on Psilocvbe annulate arrangement. Veil developed in young stages, as white, thin subarachnoid membrane, which sometimes forms an ephemeral subannulus. Context white and fleshy in pileus, fibrous, whitish to palé brownish or blackish in stipe. Pseudorhiza well developed, as a long, thick cordon or as thick, pseudofleshy, piriform, white mass. Odor and taste farinaceous. Spore print dark brown-violaceous. Basidiospores (5-)6-7(-8) x (3-)3.5-4.5(-5) x 3-4 pm, narrowly subellipsoid, oblong ellipsoid or obscurely subrhomboid in face-view, subellipsoid in side-view, thin-walled, wall 0.5-0.8 ¡am thick, palé to dark yellowish-brown, with a trún- cate germ pore, and a short and acute apiculus. Basidia (13-)(15-)(20-)22-29 x (4-) (5-) 6-7 pin, 4-spored, clávate or subvesiculose-subcylindric, with a middle con- striction, hyaline. Pleurocystidia (12-)15-20(-24)(-28) x (3-)4-6(-8)(-10)(-12) pm, common, but difficult to find, hyaline, bottle shaped, subfusiform, subcylindric or subventricose, with a wide or narrow base, mucronate or with a short to very pm long neck up to 28 long., sometimes sublageniform, irregularly branched. Pseudocystidia (18-)(21-)25-33 (-40)(-56) x (6-)8-10(-16)(-17.5) pm, common, gray- ish, polymorphous, subfusiform, subventricose, subglobose or sublageniform, branched or lobulated, sometimes submoniliform, with a narrow or wide base, which is born in the trama hymenphoral. Cheilocystidia (14-)20-30(-40) x (4-)5- 7(-8)(-10)(-20) pm, generally polymorphous, sublageniform or lageniform, regular or irregularly branched or lobulate, subcylindric or globose to subglobose, with a short or long neck, with a wide or narrow base, hyaline. Pileipellis subgelati- pm pm nous, thin to thick, up to 15(-20) thick, hyphae 1.5-5 wide, thin-walled, hyaline to yellowish. Pileocystidia 16-40 x 6-8(-10) pm, ventricose-subcylindric, submoniliform or subglobose, some strangled. Pileus trama with hyphae (1.5-)3-8 pm wide, some globose, up to 30 pm wide, thin-walled, hyaline to brownish, not incrusted. Subhymenium subcellular, elements 2-5 pm wide, hyaline and incrust- ed with yellowish-brown pigment. Trama hymenophoral regular or subregular, hyphae 2-20(-26) pm wide, thin- or thick-walled, some of them inflated, hyaline, pm occasionally incrusted with yellowish-brown pigment. Oleiferous hyphae 4-7 wide, infrequent, grayish or yellowish-gray. Caulocystidia (8-)ll-25(-30)(-40) x (3.5-)7-8(-10) pm, polymorphous, sublageniform, ventricose-rostrate, subglobose orpyramidal, sometimes regular or irregularly branched, hyaline, solitary or in small groups. Clamp connections present. This new emendation is supported mainly by the description of the pleuro- cystidia and pseudocystidia, as well as more details on the cheilocystidia, pileipellis and caulocystidia, and on the size and color ofthe basidiome, and the pseudorhiza. Concerning the distribution ofP. zapotecorum it is common in the Neotropics from , 85 Acta Botánica Mexicana 100 79-106 2012 : ( ) m México to Argentina, through mountainous cloud forests, at 900-2000(-3500) el- evation in the north, or in subtropical forests in fíat lands at sea level in the south (e.g. Brazil and Argentina). There are records ofP. zapotecorum from México, Gua- temala, Colombia, Brazil, Venezuela, Ecuador, Perú and Argentina. There are evidences of the traditional use of this mushroom and taxonomi- cally cióse species (e.g. P. moseri and P. muliercula from prehispanic times. These ) inelude pieces by ceramic, stone and metal from México, Colombia and Perú. It is possible to find in these pieces, figures resembling P. zapotecorum and closely allied species, as will be discused below. Moreover, in some códices at México, such as the Magliabechiano (Fig. 32), there are possible representations ofP. zapotecorum , P. muliercula and also P. caerulescens (Figs. 1, 3, 4, 9, 10). These two latter species have basidiome very similar to that ofP. zapotecorum. As the latter has the widest distribution of the three, P. zapotecorum is considered the mushroom most repre- sented on these anthropological figures and códices. At present, P. zapotecorum together with other species such as P. caerulescens P. muliercula, P. hoogshagenii , and P. mexicana are used in nocturnal ceremonies only in México, among several ethnic groups. Studying some anthropological ceramics pieces found in México, especially from the Capacha culture in the State of Colima, in the Nevado de Colima región, Figs. 9-10. Two importantethnomycological speciesofPsilocybeinMéxico. 9. P. zapotecorum (by Halling); 10. P. muliercula (by Soria). Scale bar: 9 = 20 mm, 10 = lOmm 86 Guzmán: New taxonomical and ethnomycological observations on Psilocybe Figs. 11-18. Microscopic features oíPsilocybe mairei andF! zapotecorum. 11-13. P. mairei : P 11. basidiospores, 12. basidia, 13. cheilocystidia. 14-18. zapotecorum 14. basidiospores, : 15. basidia, 16. pleurocystidia, 17. pseudocystidia, 18. cheilocystidia (all from the holotype). Scale bar: 4 pm. 87 Acta Botánica Mexicana 100 79-106 2012 : ( ) an interesting piece (Fig. 19) was found and presented for first time by Furst (1974) and later by Schultes and Hofmann (1979). It is a piece of about 15 cm tall and de- posited in a private collection (Furst, 1974). They described this piece as “mushroom ceremony” and “dancing Indians”, respectively, around a long-stemmed mushroom effigy, which was suggested to be a Psilocybe by Furst (1974) or P. mexicana or a closely related species by Schultes and Hofmann (1979). However, due to the ro- bustness of the mushroom and the thick stipe, it is probable that the species is P. zapotecorum (Figs. 1, 2, 9), a mushroom common in Colima región. The four per- sonages of this piece are embracing each other, and are not really dancing. This is because their faces with their eyes out oforbit suggest that they are under the effects ofneurotropic mushrooms. This is the reason that they need to embrace themselves, because under the influence ofneurotropic fungi, a person can neither remain stand- ing ñor dancing. Also, in this figure we can see the effects ofgigantism or dwarfism that are commonly produced by the neurotropic mushrooms. However, a most important observation on this Fig. 19 which had not been noted until now either by anthropologists or other specialists, is that the headdress or hat (much like a turban of the Oriental people) of the four personages is really a snake, as too are the arms. This coincides with the fact that snakes were considered sacred animáis and thus very important in the religión of several Mexican Indian cultures, such as the Náhuatl (or Aztec). For the Nahuatls, snakes represented the god Quetzalcóatl. This is also so for the Teotihuacán culture, where the representa- tion of Quetzalcóatl is covered by many large stone snakes heads. Also Schultes (1939, 1940) observed that Quetzalcóatl is represented with mushrooms in several figures in the Vindobonensis Codex from the Mixtee culture (in Oaxaca), as dis- cussed later by Wasson (1980). This relationship with Quetzalcóatl is possible to confirm in another figure from the same Capacha culture, found also in the Nevado de Colima región (Fig. 20). In this latter, five embraced Indians form a circle around a central personage similar to them, with the same snake hat, but also having arms as snakes, are hands as the snake heads. This piece was reported by Donitz et al. (2001) from the Museo Universitario de Arqueología at Manzanillo, but without any com- ments. Note that both figures (19 and 20) are very similar between them, with only one major difference, i.e., that in the second there is not any mushroom but instead at the center is an important personage with snakes, who is suggested to be Quetzal- cóatl. Thus with this latter figure together with the other, it is possible to conclude that the ingestión of sacred mushrooms like psilocybes, is probably related to the cult of Quetzalcóatl. If these two figures are indeed related with Quetzalcóatl, this means that the Capacha culture was probable under the influence ofthe Náhuatl (the 88