"Ann. Mus. civ. Rovereto
Sez.: Arch., St., Sc. nat.
Vol. 14 (1998) 189-280
2000
GASTÓN GUZMÁN, JOHN W. ALLEN & JOCHEN GARTZ
A WORLDWIDE GEOGRAPHICAL DISTRIBUTION OF THE NEUROTROPIC FUNGI, AN ANALYSIS AND DISCUSSION
Abstract - GASTÓN GUZMÁN, JOHN W. ALLEN & JOCHEN GARTZ - A worldwide geographical distribution of the neurotropic fungi, an analysis and discussion. In this paper, the world distribution of 216 known species of neurotropic fungi is discussed. The neurotropic fungi considered are divided into the following four groups: 1) species with psilocybin and related indoles, or those likely to contain these substances, 2) species with ibotenic acid, 3) ergot fungi, and 4) species used as sacred fungi for which no reliable chemical studies have been found. In the first group are Psilocybe (116 species), Gymnopilus (14 species), Panaeolus (13 species), Copelandia (12 species), Hypholoma (6 species), Pluteus (6 species), Inocybe (6 species), Conocybe (4 species), Panaeolina (4 species), Gerronema (2 species) and Agrocybe, Galerina and Mycena (each with one species), although in several species of this group, mainly in the Panaeoloideous fungi, there are no known chemical studies. In the second group are Amanita muscaria, A. pantherina and A. regalis; in the third group are Claviceps purpurea and allies: 5 species of Claviceps and 2 of Cordyceps; in the fourth group are bolets (two genera with 8 species), Russula (6 species), and 6 species of gasteromycetes (Lycoperdales y Phallales) in 3 genera. Concerning the distribution of Psilocybe, the majority of the species are found in or near the Austral hemisphere, mainly in the subtropical humid forests. Within these forests reside the most most well documented ethnic groups that use neurotropic fungi, such as the native peoples of Mexico and New Guinea. Mexico has the highest number of neurotropic species of fungi, with 76 species, of which 44 belong to Psilocybe (39 % of the world). More than 450 bibliographic references were considered. Keywords: Neurotropic fungi, Check list, Distribution Riassunto - GASTÓN GUZMÁN, JOHN W. ALLEN & JOCHEN GARTZ - Distribuzione mondiale dei funghi psicotropi: analisi e discussione. Nel presente articolo viene discussa la distribuzione mondiale delle 216 specie sinora note di funghi psicotropi. Questi funghi sono suddivisi in quattro gruppi: 1) specie contenenti psilocibina e affini composti indolici, o probabilmente contenenti questi composti; 2) specie contenenti acido ibotenico; 3) specie di ergot; 4) specie usate come funghi sacri per le quali non sono
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disponibili studi chimici certi. Al primo gruppo appartengono membri dei generi Psilocybe (116 specie), Gymnopilus (14), Panaeolus (13), Copelandia (12), Hypholoma (6), Gerronema (2) e Agrocybe, Galerina e Mycena (ciascuna con una specie), sebbene per numerose specie di questo gruppo, principalmente funghi Panaeoloidei, non siano noti studi chimici. Al secondo gruppo appartengono Amanita muscaria, A. pantherina e A. regalis; al terzo gruppo appartengono Claviceps purpurea e specie affini: 5 specie di Claviceps e 2 di Cordyceps; nel quarto gruppo rientrano boleti (due generi con 8 specie), 6 specie di Russula e 6 specie di Gasteromycetes (Lycoperdales e Phallales) distribuite in 3 generi. Riguardo alla distribuzione delle Psilocybe, la maggior parte delle specie si trova nel o vicino allemisfero Australe, principalmente nelle foreste umide subtropicali. Allinterno di queste foreste risiedono i gruppi etnici più importanti che usano i funghi psicotropi, quali le popolazioni native del Messico e della Nuova Guinea. Il Messico possiede il numero più elevato di specie neurotropiche di funghi, con 76 specie, delle quali 44 appartengono al genere Psilocybe (39% delle specie nel mondo). Vengono considerati più di 450 riferimenti bibliografici. Parole chiave: Funghi psicotropi, Check list, Distribuzione
INTRODUCTION The fungi with neurotropic (hallucinogenic or psychotropic) properties, also referred to as hallucinogenic, narcotic, magic, sacred, psychedelic or entheogenic mushrooms, are highly diverse and have a wide distribution throughout the world. During the past 40 years, since the rediscovery of the traditional use of hallucinogenic fungi in Mexico among several groups of indigenous peoples native to the central or southern regions of the country, numerous species of neurotropic mushrooms have been identified (Guzmán, 1959, 1977a, 1978b, 1982, 1983, 1990a, b). They were first studied by Schultes (1939), Singer (1949, 1958, 1959, 1960a), Singer and Smith (1958), Heim (1956a, b, 1957a, c, 1958a, b), Wasson and Wasson (1957), Heim & Wasson (1958) and Wasson (1959a, b, 1962, 1980). These fungi were so important in the traditions of Mexico, that Guzmán (1997) reported more than two hundred common names of them, many in Indian languages, as «apipiltzin», «atkad», «di nizé taaya», «shi thó», and «teotlaquilnanácatl» (which translate to: kid or little boy, mayor or leader, fungus of the genius, that eruptions thing, and divine fungus, respectively), including the unusual and rare word «teonanácatl» (divine mushroom), first reported by Sahagún (15691582) and then by Schultes (1939), which is now so indiscriminately used to name any Mexican hallucinogenic fungi. Among the most common Spanish names used by the Indians when referring to the sacred mushrooms, are «san isidros», «pajaritos» and «derrumbes» (a Spanish saint of the agriculture, little birds, and landslides, respectively). These are the names most commonly used to describe Psilocybe cubensis (1), P. mexicana and P. zapotecorum, respectively
(1) For the authors, synonyms and classification of the species see Table I, except for species not considered there.
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(Guzmán, 1959, 1983, 1997; Allen, 1997a). The studies on the neurotropic fungi in Mexico were so important, that Guzmán (1990a, b) divided the development of the Mexican mycology into the following two important time periods: 1) before Wasson, Heim and Singers studies on the hallucinogenic fungi, which had been developed between 1954-1958, and 2) after Wasson, Heim and Singers studies. This came about because the studies by these specialists involved in the neurotropic fungi played such an important role toward the study of other fungi, that they produced interest in other specialists to study all of the fungi in the country. In the late fiftys, only around 20 species of the neurotropic fungi were recognized, belonging to the genera Psilocybe (the majority), Conocybe (e.g. C. siliginoides), Stropharia (e.g. S. cubensis, later identified as Psilocybe cubensis), Panaeolus (as P. sphinctrinus), Cordyceps (two species), Claviceps (C. purpurea) and Amanita (A. muscaria); also considered were the edible species of Clavariadelphus and Gomphus, which were erroneously mixed with the properties of Cordyceps spp. (Heim & Wasson, 1958; Singer & Smith, 1958; Guzmán, 1959). Wasson and Wasson (1957) brought attention to the fact that Amanita muscaria was an important sacred fungus in the Siberian region (Russia) between the Chukchee and Koryak peoples, as were the psilocybian fungi important between the Indians in Mexico.Later, Singer (1958, 1960a), Heim and Wasson (1965) and Heim (1965a, 1978) reported the use of Psilocybe, Russula and Boletaceous fungi as sacred mushrooms among several aborigines in New Guinea. The criteria used to define the various neurotropic fungi are often confusing according to the mycologists. For example, Oldridge et al. (1989) considered some polypores known to contain hordenine, N-methyltyramine and tyramine, as psychotropic fungi, e.g. Laetiporus sulphureus (Bull.: Fr.) Murrill and Meripilus giganteus (Pers.: Fr.) P. Karst. They produce certain chemical reactions in the central nervous system which resulted in dizziness and disorientation. However, the first species is a common and important edible fungus in Mexico (Guzmán, 1977a, 1997) and in other parts of the world (Dickinson & Lucas, 1979; Metzler et al., 1992), and from the latter species there are no reports concerning its use, as there are regarding other polypores. Thoen (1982) commented on the use in several regions of some polypores in religious ceremonies for magic activities, such as Polyporus tuberaster Jacq.: Fr., Poria cocos (Schwein.) Wolf, Ganoderma lucidum (M.A. Curt.: Fr.) P. Karst., Fomes fomentarius (L.: Fr.) Kickx and others. Guzmán et al. (1975) reported the cult of Ganoderma lobatum (Schwein.) G.F. Atk. in a church in Mexico (in Chignahuapan, Puebla); that church was built especially in honor of the fungus. The Indians who reside in the region regard the fungus as a saint. This interesting fungus was found in the last century and is decorated in its inner surface with an arresting sketch, por191
traying a Christ with a sun and moon on each side of him. However, there is no evidence of neurotropic properties related to the use of this fungus, its use in the cult is probably in relationship with the use of neurotropic species of Psilocybe, used in this region where active species of this genera are commonly found. Ott (1993) presented a list of fungi species containing psilocybin based on bibliographic references. These fungi belong to the genera Agrocybe (one species), Conocybe (four), Copelandia (six), Galerina (one), Gerronema (two), Gymnopilus (seven), Hygrocybe (one), Inocybe (seven), Mycena (one), Panaeolina (two), Panaeolus (nine), Pluteus (five), Psathyrella (two), and Psilocybe (fortyseven). For each of the species of interest in these genera, Ott presented the bibliographic references about related studies, notes, as well as problems or contradictions, such as those in Gerronema, Hygrocybe and Inocybe, according to the work of Gartz (1986e). In those bolets reported by Heim (1963, 1966, 1978) as hallucinogenic in New Guinea, Ott (1993) stated that they do not possess any neurotropic properties. Phellinus igniarus (Fr.) Quél. and Fomes fomentarius were observed in Alaska as narcotic fungi. In Gymnopilus, there is a interesting study on G. penetrans (Fr.: Fr.) Murrill (Dangy-Cave & Arpin, 1974), although apparently independent of the neurotropic fungi. Hatfield (1979) reported that ibotenic acid was present in Amanita pantherina and A. cothurnata, in spite of the reports of intoxication induced by these fungi in central Europe. Adewusi et al. (1993) considered Chlorophyllum molybdites (Meyer: Fr.) Massee from Africa with to have some neurotropic properties, based in their experiments in weanling rats and related it with the common name in the Yoruba tribe: «a jegba ariwo-orun» (meaning: «eat and hear voices from heaven»); however, many reports (Lincoff, 1981; Guzmán, 1977a; Portugal et al., 1992; Duffy and Vergeer, 1977) on this fungus considered it to be a poisonous mushroom species. Pegler (1977, 1983) commented that there is a considerable amount of confusion about wheter this species is toxic or edible. Singer (1969) said: «apparently not all forms or races are poisonous», and he reportedcases of poisoning in the U.S.A., Argentina, Phillipins and East Africa. However, Heim (1978) considered Ch. molybdites as an edible fungus in Africa. Schizophyllum commune was reported as a hallucinogenic fungus in Australia (Southcott, 1974); yet this species is a common edible fungus sold in popular markets in Guatemala and southeastern Mexico. The confusion originated because the Mazatec Indians of Oaxaca (Mexico) often referred to this fungus as «nise» (meaning: «little bird»), a name also used for Psilocybe mexicana, but without any relationship between them with respect to their properties (Guzmán, 1997). Regarding the puffballs (Gasteromycetes, Lycoperdales), Burk (1983) discussed the magic and religious uses of several unidentified species of puffballs among certain North American Indians peoples. The fungi which typically grew in circles (fairy rings) on the prairies, were referred to as «fallen stars». Guzmán 192
(1994a, 1994b, 1997) discussed several puffballs used by the Mexican Indians in traditional medicine, some of them, such as Lycoperdon perlatum Pers., forming fairy rings in grasslands. Although none of these species have neurotropic properties, they are, on the cotrary, edible. However, Heim and Wasson (1962) and Heim et al. (1967) reported the use of Lycoperdon mixtecorum and L. marginatum (synonyms of Vascellum qudenii and Lycoperdon candidum, respectively [following Guzmán, in Ott et al., 1975)] as a narcotic fungi among the Mixtec Indians of Oaxaca, Mexico. These fungi were later studied by Ott et al. (1975) in the same locality where Heim and Wasson (1962) first found them. Ott et al. (1975) observed that Heim and Wassons fungi are edible and common in Mexico as reported by Guzmán (1977a, 1997), but in the Mixtec zone they are used in a manner suggesting the confusion of these species with more traditional neurotropic fungi among some Indians for religious or magical proposes. Besides these two fungi, Ott et al. (1975) identified yet another six «sacred» species of fungi from the same locality, as Vascellum pratense, V. curtisii (Berk.) Kreisel, V. intermedium A.H. Sm., Lycoperdon oblongiosporum, Rhizopogon sp. and Astraeus hygrometricus (Pers.) Morg., reported that the Indians used them indistinctly as a narcotic fungi, along with Scleroderma verrucosum Pers., which was experimentally proven to be poisionous. Chemical analysis of these fungi (except inS. verrucosum) confirmed the presence of psilocybin. The conclusions of Ott et al. (1975) were that the Mixtecnarcotic puffballs were a mixture of at least nine species of fungi containing no neurotropic properties. However, Schultes and Hofmann (1973, 1979) considered Heim and Wassons fungi among the «narcotic fungi». In spite of the above observations these fungi are considered in the present work due to their popularity between the Indians and are listed in the bibliography (e.g. Schultes, 1976). Another example with similar discrepancies among various authors and their resulting conclusions is Dictyophora indusiata, with its three forms (Guzmán et al., 1990) (see Table I), all of which are used as special «narcotic» fungi for divination purposes among the Chinantec Indians in Oaxaca, Mexico (Heim and Wasson, 1958; Wasson, 1959a; Guzmán, 1997). Recently, chemical studies on species of neurotropic fungi show the presence of psilocybin, related compounds mistaken for psilocybin, or indole metabolites in several fungi (Becker et al., 1988; Besl, 1994; Chilton, 1978; Christiansen et al., 1984; Gartz, 1985a, b, c, 1986a, b, d, 1987a, c, 1989a, b, c, e, 1991a, b, 1995a; Gurevich, 1993; Koike et al., 1981; Kreisel and Lindequest, 1988; Semerdzieva et al., 1986; Singer, 1978; Stijve, 1987; Stijve and Bonnard, 1986; Stijve et al., 1985 and Takemoto et al., 1964a, b, c). However, several studies must be considered doubtful due to errors in analysis, as pointed out by Ott (1993) and Stijve (1995). Bresinsky and Besl (1990) considered those studies on the hallucinogenic principles of Stropharia cyanea (Bolt. ex Secr.) Tuomikoski [also known as Psilocybe caerulea (Kreisel) Noordeloos] and 193
Stropharia caerulea Kreisel [Psilocybe caerulea (Kreisel) Noordeloos] (Noordeloos, 1995), S. coronilla (Bull.: Fr.) Quél., Mycena pura (Pers.: Fr.) P. Kumm. andAmanita gemmata (Fr.) Bertillan to be doubtful. Samorini (1989) pointed out the same with Mycena pura. Stijve and Kuyper (1988) did not find psilocybin in Psathyrella candolleana (Fr.) Maire, Rickenella swartzii (Fr.) Kuyp., Gerronema fibula, Gymnopilus fulgens, G. spectabilis, Hygrocybe psittacina (Fr.) P. Karst. and H. psittaccina var. californica Hesler & A.H. Sm. Stivje and Meijer (1993) failed to find psilocybin and other psilocybian compounds in Gymnopilus spp., Panaeolina foenisecii and Rickenella straminea (Petch) Pegler. Frequently, a single species has been reported with and without neurotropic substances according to different researchers. An example is Panaeolina foenisecii, a very common fungus in the prairies of many parts of the world. Mantle and Waight (1969), Ott and Guzmán (1976), Beug and Bigwood (1982), Stijve (1987) and Stijve et al. (1984) did not find any psilocybin or psilocin in this species, but other papers, e.g. Robbers et al. (1969), Olah (1969), Fiussello and Ceruti-Scurti (1972), Pollock (1976) and Bresinsky and Besl (1990) reported psilocybin. Allen and Merlin (1992c) discussed doubts on the psychoactive properties of this fungus. It seems that P. foenisecii is a toxic fungus, more so than neurotropic, as is with the majority of the Panaeolodeous fungi. Regarding Conocybe siligineoides, a species reported by Heim (1956b) and Heim and Wasson (1958) as a sacred mushroom in Mexico, no chemical studies have been made on this species to date, but C. cyanopus and C. smithii were shown to contain psilocybin (Benedict et al., 1967) (Mantle and Waight, 1969, wrote erroneously that Benedict et al., 1967, reported C. siligineoides contained psilocybin). It is important to observe that C. siligineoides was collected only one time in 1955 by Wasson in the State of Oaxaca, Mexico, and there have been no additional reports of this species being found since. Even after several years of extensive field-work in Mexico, Guzmán has been unable to re-collect this fungus (Guzmán, 1997). The problem of mis-identification is yet another factor contributing to the confusion concerning published chemical studies of neurotropic fungi. Unfortunately, it is frequently found that many chemical studies do not have a taxonomic base, in some cases a mixture of different species were studied. Guzmán, found a mixture of Panaeolus spp. and Psilocybe mexicana, together with P. coprophila (Bull.: Fr.) P. Kumm., all of them identified as P. coprophila (in ENCB Herbarium at Mexico City), in material used by Leslie and Repke to isolate psilocybin (Guzmán, 1983). Psilocybe pseudobullacea (Petch) Pegler is a not bluing species (Guzmán, 1983, 1996) and no neurotropic properties have been found. However, Marcano et al. (1994) isolated psilocybin and psilocin from Venezuelan specimens; it is probable that the Venezuelan material belongs to a neurotropic species that has yet to be determined as such. Høiland (1978) reported psilocybin in P. atrobrunnea. It is probable that Høilands fungus is a species 194
with similar appearance, such as P. coprinifacies or P. maire, since P. atrobrunnea is not a neurotropic fungus (Guzmán, 1983). In the chemical studies on fungi, the age of the studied specimens is an important variable which needs to be taken into consideration. Repke et al. (1977a) showed variations in the presence of baeocystin according to the age of the studied materials, e.g., in Psilocybe baeocystis and P. cyanescens have no trace of indoles in specimens analysed 20-60 days after the collection. This explains why the Mexican Indians wisely say in relationship with the use of the sacred mushrooms, that the old dried specimens kept for more than one yearare not good to use, and they throw them out. One of the authors (Guzmán), observed in an experiment that normal doses of hallucinogenic fungi (Psilocybe mexicana in one case, and P. caerulescens in other), were only slightly neurotropic or entirely inactive in the persons who ate them, because the fungi were kept dried for almost a year. Ohenoja et al. (1987) detected a decrease of psilocybin in herbarium specimens of P. semilanceata, according to the age of the collections. They found 0.014, 0.67, and 0.84 % dry wt. in specimens from 1869, 1954 and 1976, respectively. It seems that psilocybin and psilocin are volatiles, as Guzmán observed while exploring Oaxaca (Mexico) looking for neurotropic fungi in 1958.He experienced colored hallucinations although he had not consumed any fungi.This occurred one night when he was trying to sleep in a small closeddoor..."
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