An Overview on the Taxonomy, Phylogenetics and Ecology of the Psychedelic Genera Psilocybe, Panaeolus, Pluteus and Gymnopilus
Dominique Strauss
Soumya Ghosh
Zurika Murray
Marieka Gryzenhout- Department of Genetics, Faculty of Natural and Agricultural Sciences, University of the Free State, Bloemfontein, South Africa
Psilocybin and psilocin, two psychoactive components found in “magic mushrooms,” have therapeutic potential in a number of mental health disorders without the addictiveness and overdose risks found in other mind-altering drugs, such as cocaine, methamphetamines and alcohol. Psychedelic mushrooms occur naturally, are wide distributed and easily accessible. The need for reviews and comprehensive field guides is urgent due to the recent surge of research into psychedelic mushrooms along with public interest. Psilocybin and psilocin are recorded in mushroom species of Psilocybe, Panaeolus, Pluteus, and Gymnopilus. This review discusses species identification, taxonomy and classification, available DNA sequence data and psychedelic species in Psilocybe, Panaeolus, Pluteus, and Gymnopilus, as well as similar looking genera that could be harmful.
Introduction
Over 30 million people have used psychedelic substances at least once throughout their life (Krebs and Johansen, 2013). Psychedelic mushrooms, also referred to as hallucinogenic mushrooms (Nichols, 2016), are the most widely used natural hallucinogen in the world due to their wide geographical distribution and easy cultivation (Stafford, 2013). Their existence played a substantial role in defining the psychedelic era of the 1960s and 1970s, and psychedelic cultural customs remain in places where “magic mushrooms” remain legal, such as the Bahamas, Brazil, British Virgin Islands, Jamaica, Nepal, Netherlands, and Samoa (Flaherty et al., 2017; Duneman, 2021).
Many states in the United States of America have started making changes in legislations regarding the use and culturing of psychedelic mushrooms (Hartman, 2018). Popular intellectuals and entertainers advocate the positive uses of psychedelics to millions of people using podcasts and virtual videos (Kohn and Hofmann, 2010; Hartman, 2018; Rogan, 2018, 2019). Growers’ guides are available (Oss and Oeric, 1991) and billionaires are also investing heavily into psychedelic research (Marlan, 2019).
Psilocybin, a serotonergic hallucinogen, is the main psychoactive substance found in psychedelic mushrooms, and alters perception and mood, and produces hallucinations in individuals who ingest them (Nichols, 2016). To date, human and animal studies have shown that psilocybin is non-addictive and has short- and long-term benefits in mood disorders, abuse disorders and chronic pain (Amsterdam et al., 2011; Carhart-Harris et al., 2016; Hanks and González-Maeso, 2016; Tyls et al., 2016; Hartman, 2018; Johnson et al., 2018; Dos Santos et al., 2019; Castellanos et al., 2020; Reiff et al., 2020).
Psychedelic drug research will increase globally and so will the research on at least 300 species of psychedelic mushrooms, divided across Psilocybe, Panaeolus, Pluteus, Gymnopilus and other genera also containing psychedelic species such as Amanita, Copelandia, Inocybe, and Pholiotina (Guzmán et al., 1998; Metzner, 2005; Reingardiene et al., 2005; Oxford Analytica, 2021). This review provides an overview of only Psilocybe, Panaeolus, Pluteus, and Gymnopilus, their current taxonomy and ecology. Brief molecular data for species in each genus is also provided. For any party involved in research or products of research on these fungi and their metabolites and the public, identifying these often similarly looking, inconspicuous mushrooms are a challenge. The current review of these psychedelic mushrooms would be of great value to researchers, civilian scientists, medical professionals, mushroom hunters as well as the public.
Identification of Psychedelic Mushrooms
Most professional and citizen mycologists rely on morphology for identification of mushrooms based on characteristics of the cap, stem, gills, spores, spore-bearing structures and habitats of mushrooms (Goldman and Gryzenhout, 2019). Psychedelic mushrooms are often small, indistinct, brown to white mushrooms that, usually but not always, bruise bluish to black when the tissue is cut or damaged (Guzmán, 2008). The blue stains result from an oxidative reaction when psilocybin is exposed to oxygen (Lenz et al., 2020).
Regional field guides and checklists are regularly consulted for more identification characteristics but, few dedicated guides exist to aid in the identification of psychedelic mushrooms (Gartz, 1996; Stamets, 1996). Field guides tend to use generalised descriptions of morphological characteristics which are inefficient to cover the more than 300 species of psychedelic mushrooms currently known (Stamets, 1996). For example, in South Africa, few psychedelics have been described in previous field guides and checklists (Kinge et al., 2020), with better inclusion only achieved recently (Van der Walt et al., 2020; Gryzenhout, 2021).
An additional hurdle making psychedelic mushroom identification difficult is the continuous variation in cap, stem and gill morphology (Guzmán, 2009). Expert scientific publications must frequently be consulted for the use of microscopic features. These resources are not always available or difficult to find for members of the public or others studying these mushrooms, such as law enforcement officers and medical researchers. Although available studies include increasingly more detailed descriptions of species, there is currently no up to date monograph or guide (Høiland, 1978; Guzmán et al., 2004, 2016; Borovička, 2008; Ramírez-Cruz et al., 2013a; Cortés-Pérez et al., 2021).
DNA technologies are allowing researchers to identify fungi at faster rates and higher accuracy than morphological methods (Mullineux and Hausner, 2009; Zhang et al., 2016; Badotti et al., 2017; O’Hanlon, 2018). The combination of morphological and molecular methods of identification have been used to split genera, such as was the case for Psilocybe and Deconica, and transfer species, previously thought to be psychedelic, to non-psychedelic genera (Moncalvo et al., 2002; Matheny et al., 2006; Norvell et al., 2010; Ramírez-Cruz et al., 2012). Molecular methods have been used to distinguish over 100 species in Psilocybe, Panaeolus, Pluteus, and Gymnopilus (Ma et al., 2014; Menolli et al., 2014; Khan et al., 2017; Malysheva et al., 2019).
The most frequently sequenced region of DNA used for these genera is the Internal Transcribed Spacer (ITS) region (approximately 500–800 bp), which forms a part of the ribosomal operon (Schoch et al., 2012; Badotti et al., 2017). The ITS region is used due to its efficacy at identification across fungal lineages, and represents the primary barcoding region (Schoch et al., 2012). The region is sufficiently variable to allow single nucleotide polymorphisms or more to differentiate one species from another. Wesselink (2018) used this region to convey its accuracy identification of psychedelic mushrooms in forensic settings. Other gene regions have also been used, including the translational elongation factor 1α (TEF1α), RNA polymerase II (rpb), large subunit ribosomal ribonucleic acid (nLSU) and small subunit ribosomal ribonucleic acid (nSSU) (Tóth et al., 2013; Meyer et al., 2019; Tekpinar and Kalmer, 2019). However, sequences of all the genes are not available for most species.
The Most Important Genera Containing Psychedelic Species
Psilocybe
Psilocybe sensu stricto is most likely the best known genus containing over 150 species distributed worldwide and represented by the type species Psilocybe semilanceata (Figure 1A; Redhead et al., 2007; Norvell et al., 2010). Mushrooms are described as little brown mushrooms (LBMs) or little white mushrooms (LWMs) (Li et al., 2014; Dalefield, 2017) with a viscid cap when moist, an dark to purplish black coloured spores (Arora, 1986) and a dark purple-brown spore print (Estrada et al., 2020). Other characteristics include a separable gelatinous pellicle, fringed whitish gill edges, and typically collyboid or mycenoid aspects (Stamets, 1996; Ramírez-Cruz et al., 2013b). The caps and stems of some species may bruise a greenish-blue, similar to that of a human bruise, when the mushroom is damaged, aged or drying indicating the presence of psychedelic compounds (Lenz et al., 2020). Psilocybe are regularly found in substrates such as soil, dung, wood, and mosses (Stamets, 1996).
Figure 1. (A) Psilocybe semilanceata, (B) Psilocybe chuxiongensis, (C) psychedelic Psilocybe stuntzii (white arrow), growing across a deadly poisonous mushroom, Galerina autumnalis (black arrow) (Stamets, 1996), (D) Panaeolus cyanescens, (E) the most novel species, Panaeolus axfordii, (F) unidentified Psathyrella, (G) Pluteus cervinus, (H) Pluteus cyanopus, and (I) Pluteus salicinus, showing bluing (white arrow), (J) Gymnopilus liquiritiae, (K) Armillaria mellea, morphologically similar to G. spectabilis (L). Images obtained from Mushroom Observer (2021).
Guzmán (1983) and Singer (1986) recognised Psilocybe in the Hymenogastraceae, and included non-bluing and non-psychedelic species (Redhead and Guzmán, 1985; Stamets, 1996; Guzmán, 2012). Moncalvo et al. (2002) showed the genus to be polyphyletic with psychedelic species grouping the Strophariaceae and non-psychedelic species grouping the Hymenogastraceae (Moncalvo et al., 2002; Matheny et al., 2006). Redhead et al. (2007) proposed to split the genus and conserve the name Psilocybe for psychedelic species, represented by the type species Psilocybe semilanceata (Figure 1A), and transfer the remaining non-psychedelic species to Deconica, typified by Deconica physaloides (Ramírez-Cruz et al., 2012). The proposal was accepted unanimously by the Nomenclature Committee for Fungi (Norvell et al., 2010). Since then more species have been transferred to Deconica and multiple gene regions have supported the positions of the two genera, including nLSU-rRNA, 5.8S rRNA, and rpb1 (Ramírez-Cruz et al., 2013b).
Internal classification of Psilocybe s. str. is largely based morphologically (Maruyama et al., 2006; Ramírez-Cruz et al., 2013b; Borovička et al., 2015). Three classifications have been proposed for the division of Psilocybe into subgenera (Guzmán, 1978; Singer, 1986; Noordeloos, 2011). Psilocybe was divided into 16 subgenera by Guzmán (1978), according to the form and structure of fruiting bodies, form and wall thickness of the spores, and absence or presence of pleuro- and cheilocystidia. Singer (1986) placed all bluing psychedelic species in Caerulescentes, and distinguished six subgenera. Noordeloos (2011) assessed species on the presence of chrysocystidia, bluing and DNA sequences of the ITS, partial nLSU rDNA, and rpb1 genes, and recognised a total of three unnamed sections, that did not support previous subgenera.
Morphological characteristics of Psilocybe species often include bluing, indicative of the presence of psilocybin, for example, recently described in Psilocybe chuxiongensis, Psi. cinnamomea, Psi. thaiaerugineomaculans, Psi. thaicordispora, Psi. thaiduplicatocystidiata, and Psi. thaizapoteca (Guzmán et al., 2012; Li et al., 2014; Ma et al., 2014). Psilocybin can be accurately measured in bluing species using physico-chemical methods (Passie et al., 2002). Thirty-three and 55 species of Psilocybe have been shown to be positive for psilocybin using physico-chemical methods (Wurst et al., 2002; Andersson et al., 2009). Allen (2012) reported 146 species that either had bluing reactions or confirmed the presence of psilocybin and psilocin using physico-chemical methods.
Phylogenetics of gene sequences have described the newest species of Psilocybe. Li et al. (2014) and Ma et al. (2014) described Psilocybe chuxiongensis (Figure 1B) and Psilocybe cinnamomea from China based on the ITS, nrLSU, rpb2, and tef1-α genes. In Thailand, Psi. thaiduplicatocystidiata and Psi. thaiaerugineomaculans were described using the ITS and nLSU sequences (Ma et al., 2016), similarly to Psi. angulospora from Taiwan (Wang and Tzean, 2015). Eighty one species of Psilocybe have genome sequence data available, including Psi. tampanensis, Psi. azurescens, Psi. galindoi, Psi. cubensis, Psi. cf. subviscida, and Psi. cyanescens (McKernan et al., 2021; NCBI, 2021).
No toxic or lethal Psilocybe species are known. However, there are several poisonous species in Conocybe, Galerina, and Inocybe that could be lethal and appear morphologically similar to Psilocybe (Stamets, 1996; Figure 1C). For example, Galerina autumnalis looks similar to Psilocybe stuntzii (Figure 1C), and results in sharp abdominal pain, violent vomiting, liver and kidney failure leading to possible death in 7 to 10 days if consumed accidentally (Stamets, 1996).
Panaeolus
Panaeolus contains between 15 (He et al., 2019), 20 (Ola’h, 1969) and 29 species (Gerhardt, 1996) species worldwide and is represented by the type species Panaeolus papilionaceus (Quélet, 1872; Gerhardt, 1996). They are characterised as LBMs with small, greyish, brown or blackish, conical- or bell-shaped caps, elongated slender stalks, attached gills and a dark brown to purple-brown to black spore print (Rumack and Spoerke, 1994; Gerhardt, 1996; Stamets, 1996; He et al., 2019). The most conspicuous characteristic is the gills that become mottled with shades of grey and black while the spores mature (Rumack and Spoerke, 1994). Microscopic characteristics include spores with an apical germ pore and a cellular pileipellis (Quel, 1872; Quélet, 1872). Panaeolus are coprophilic and frequently reported in livestock droppings, including horse, cow, buffalo and elephant dung (Ediriweera et al., 2015; Wang and Tzean, 2015). Non-coprophilic species, such as Panaeolus bisporus, grow in grassy areas (Senn-Irlet et al., 1999).
The family relationships of Panaeolus remain complex. The genus was formerly recognised by Singer (1949) in the Coprinaceae, but then transferred to the Strophariaceae based on basidiospore pigmentation and the presence of chrysocystidia (Kuihner, 1980) and DNA sequences of the nLSU (Hopple and Vilgalys, 1999). He et al. (2019) and Wijayawardene et al. (2020) placed Panaeolus in incertae sedis, a taxonomic group reserved for taxons where broader relationships are unknown or undefined. However, recent DNA sequence data placed the genus in Galeropsidaceae (Kalichman et al., 2020).
At least 13 species of Panaeolus are bluing and potentially psychedelic including Panaeolus africanus, Pan. ater, Pan. castaneifolius, Pan. fimicola, Pan. microspores, Pan. moellerianus, Pan. olivaceus, Pan. papilionaceus, Pan. retirugis, Pan. rubricaulis, Pan. sphinctrinus, Pan. Subbalteatus, and Pan. venezolanus (Guzmán et al., 1998). Currently, Pan. cyanescens is the most well-known psychoactive representative (Figure 1D; Badham, 1984) and contains high levels of psilocybin, that are even higher than those found in Psilocybe species (Musshoff et al., 2000; Maruyama et al., 2003). Pan. axfordii is the most recent novel species that has also been reported to have psychedelic properties (Figure 1E; Hu et al., 2020).
The ITS region has been used in descriptions of various species, such as Panaeolus axfordii, Pan. antillarum, Pan. sphinctrinus, Pan. acuminatus, Pan. antillarum, Pan. campanulatus, Pan. retirugis, Pan. rickenii, Pan. semiovatus, Pan. alcis, Pan. bisporus, Pan. foenisecii, Pan. plantaginiformis, Pan. desertorum, Pan. papilionaceus, Pan. cambodginiensis, Pan. subbalteatus, Pan. campanulatus, and Pan. retirugis (Maruyama et al., 2006; Sette et al., 2010; Razaq et al., 2012; Osmundson et al., 2013; Ma, 2014; Ediriweera et al., 2015; Wang and Tzean, 2015; Boy et al., 2016; Malysheva et al., 2019; Hu et al., 2020). Pan. subbalteatus has a sequence available for the elongation factor subunit 1α gene region (Li et al., 2019), and Pan. acuminatus for the nLSU region (Moncalvo et al., 2002). Pan. papilionaceus and Pan. cyanescens have genome sequences available (NCBI, 2021).
Panaeolus is similar in morphology to Psathyrella, which is frequently found on wood or lignin-enriched soils and can be distinguished from Panaeolus by a brittle white stipe (Figure 1F; Kaur et al., 2014). An additional characteristic for identification involve the fading of spore colour in concentrated sulphuric acid. For example, Psathyrella present fading spores while Panaeolus spores will not be effected (Kaur et al., 2014). Panaeolina is morphologically similar but distinguished by having ornamented spores and dark brown gills, in comparison to smooth basidiospores and mottled greyish-black gills in Panaeolus (Kaur et al., 2014; Kalichman et al., 2020). There are no recorded fatal species of Panaeolus, Psathyrella or Panaeolina, but Panaeolus subbalteatus, and Panaeolus retirugishas have been reported as poisonous (Watling, 1977; Chen et al., 2014; Li et al., 2019).
Pluteus
Pluteus is a large genus of at least 500 species and is typified by Pluteus cervinus (Figure 1G; Kirk et al., 2008; Justo et al., 2011b; Wijayawardene et al., 2020). Species include small, brown- or white-capped mushrooms (1 to 15 cm across) that start conical or convex, flattening out to a typical mushroom shape with many caps having a raised central umbo (Iliffe, 2010). Furthermore, the genus is characterised by free lamellae and the absence of an annulus and volva (Menolli et al., 2010). Microscopic characteristics include smooth and round ellipsoid spores, that produce a pink spore print, and the presence of pleurocystidia and inverse hymenophoral trama (Hosen et al., 2019). Since the transfer of Chamaeota mammillatus to Pluteus using morphological and DNA sequences, the description of the genus also includes species with a partial veil (Minnis et al., 2006; Minnis, 2008; Menolli et al., 2010). Pluteus species are common in tropical habitats and grow almost exclusively on well-decayed wood (Justo et al., 2011b; Desjardin and Perry, 2018).
Pluteus is part of the Pluteaceae family (Wijayawardene et al., 2020). Morphologically the genus is divided into the subgenera Pluteus, Hispidoderma and Celluloderma (Singer, 1958; Shaffer and Singer, 1976). Subgenus Pluteus possesses metuloid pleurocystidia, a pilepellis as a cutis; Hispidoderma have non-metuloid pleurocystidia and a pileipellis composed of elongated elements organised as a cutis, and an hymeniderm or a trichoderm; and Celluloderma have non-metuloid pleurocystidia and a pileipellis composed of short, clavate or spheropedunculate elements organised as an hymeniderm, with transitions to an epithelium (Singer, 1958; Shaffer and Singer, 1976). This division of Pluteus was supported with DNA sequences of the nLSU, nSSU and ITS regions (Shaffer and Singer, 1976; Justo et al., 2011a).
Pluteus includes a few species that are psilocybin producing. These include Pluteus atricapillus, Plu. cyanopus, Plu. glaucus, Plu. nigroviridis, Plu. salicinus, and Plu. villosus (Figures 1H,I; Saupe, 1981; Guzmán et al., 1998; Allen, 2012). Psychoactive tryptamines have been found in Plu. atricapillus, Plu. glaucus, Plu. nigroviridis, and Plu. salicinus (Gartz, 1987; Wurst et al., 2002; Stríbrný et al., 2003; Andersson et al., 2009).
Many widely distributed species of Pluteus have available DNA sequence data (Rodriguez et al., 2008; Justo et al., 2014; Araujo and Sampaio-Maia, 2018; Hosen et al., 2019; Ševčíková et al., 2021). Eight species were sequenced for ITS data including Plu. brunneocrinitus, Plu. cebolinhae, Plu. crinitus, Plu. halonatus, Plu. hispidulopsis, Plu. karstedtiae, Plu. Necopinatus, and Plu. paucicystidiatus (Menolli et al., 2015) and a further 12 species were sequenced for the ITS and tef1 gene regions (Justo et al., 2014). Recently, Plu. squarrous (Hosen et al., 2019), and Plu. cervinus (Ishaq et al., 2021) have been sequenced for the ITS region. Pluteus hubregtseorum was described using both the nrITS and EF1- α genes (Ševčíková et al., 2021). One genome has been published of Pluteus cervinus (Araujo and Sampaio-Maia, 2018).
No toxic species in Pluteus are known (Halling et al., 1987). The genus is known for edible species, including Pluteus cervinus (Halling et al., 1987; Ishaq et al., 2021). Confusion with members from the Rhodophyllaceae is possible due to the Pluteaceae family sharing pink spores, attached gills, spores that are angular in outline and a regular to irregular gill trama (Halling et al., 1987).
Gymnopilus
Gymnopilus contains over 200 species world-wide and has a psychedelic type species, namely Gymnopilus liquiritiae (Figure 1J; Karsten, 1879; Wurst et al., 2002; Kalichman et al., 2020). The genus is characterised by dry, reddish-brown to rusty orange or yellow fruiting bodies that are medium to large and have a well-developed veil (Stamets, 1996). Microscopic characteristics include spores with verrucose to rugulose ornamentation, no germ pore, and dextrinoid walls, and spore prints are a rusty brown (Holec, 2005). Gymnopilus typically include lignicolous species that grow on wood as well as grassy areas with decomposing wood (Guzmán-Dávalos et al., 2003).
In traditional classifications, Gymnopilus was placed in the Cortinariaceae based on the ornamentation and lack of germinal pores of the basidiospores (Shaffer and Singer, 1976). However, based on the presence of styryl-pyrones, bis-noryangonin and hispidin the genus was transferred to the Strophariaceae (Kuihner, 1980; Rees et al., 2004; Holec, 2005). Based on DNA sequence comparisons Gymnopilus is currently characterised under the Hymenogastraceae (Kalichman et al., 2020).
Gymnopilus is subdivided into sections Annulati, which has a membranous partial veil, and Gymnopilus, which lacks a veil (Guzmán-Dávalos et al., 2003). Phylogenetic analyses based on DNA sequences does not support the subdivision (Guzmán-Dávalos et al., 2003). The ITS regions divided 38 taxa into five well supported clades, called spectabilis-imperialis, nevadensis-penetrans, aeruginosus-luteofolius, lepidotus-subearlei and an unnamed clade formed by G. underwoodii, G. validipes, and G. flavidellus (Guzmán-Dávalos et al., 2003). LSU sequence data confirmed the genus to be monophyletic (Moncalvo et al., 2002).
Novel species of Gymnopilus were characterised using the ITS region, which include G. minisporus, G. turficola, and G. dunensis (Khan et al., 2017; Liu and Bau, 2019; Bashir et al., 2020) and G. swaticus using the ITS and LSU regions (Khan et al., 2017). In a study of sequestrate fungi in the Cortinariaceae using the ITS region, three species of Gymnopilus (G. sapineus, G. penetrans, and G. spectabilis) were used as an outgroup (Peintner et al., 2001). Thomas et al. (2002) included five species of Gymnopilus (G. aeruginosus, G. penetrans, G. picreus, G. sapineus, and G. spectabilis) in a study describing Anamika, a genus of Cortinariaceae, using both the ITS and LSU gene regions. The mitochondrial genome of G. junonius, a psychedelic species, has been sequenced (Cho et al., 2021) and the genome of G. junonus and G. dilepsis is available (NCBI, 2021).
Psilocybin and psilocin are commonly found in Gymnopilus species (Andersson et al., 2009). At least 16 species of Gymnopilus are considered psychedelic, including G. cyanopalmicola, G. palmicola, G. igniculus, G. validipes, G. aeruginosus, G. braendlei, G. intermedius, G. lateritius, G. liquiritiae, G. luteoviridis, G. luteus, G. purpuratus, G. sapineus, G. spectabilis, G. subpurpuratus, G. validipes, and G. viridans (Hatfield and Valdes, 1978; Guzmán et al., 1998; Holec et al., 2003; Guzmán-Dávalos and Herrera, 2006). Other tryptamine compounds such as serotonin is also commonly found in Gymnopilus species, suggesting that the genus has species with possible undiscovered tryptamines.
Gymnopilus is morphologically similar to Armillaria, and frequently found in similar substrates, such as grassy areas and decomposing wood (Figures 1K,L; Rees et al., 1999; Spring et al., 2016). Gymnopilus spectabilis, a known psychedelic mushroom, looks alike to Armillaria mellea, an edible mushroom (Spring et al., 2016) and accidental ingestions of G. spectabilis in place of A. mellea have been recorded (Roper, 2003). Galerina is also morphology similar to Gymnopilus and a possible synonymous genus (Landry, 2016; Kalichman et al., 2020). Species of Armillaria, Galerina, and Gymnopilus (such as Gymnopilus junonius, and Gymnopilus spectabilis) (Figure 1L) have been reported as poisonous (Maeta et al., 2008; Konno, 2009; Chen et al., 2014; Lee et al., 2020).
Conclusion
As the medicinal value of psychedelic species increases, the importance of accurate species identification of psilocybin containing species runs parallel. Up to date taxonomic monographs are also needed for each of the genera containing psychoactive species. Where mostly ITS sequence data is available, more genes should be added for phylogenetics, and numerous more species remain to be sequenced. With the foundation already laid, future research on these increasingly important fungi should be easily accomplished.
Author Contributions
DS researched and wrote the manuscript. SG, ZM, and MG supervised and edited. All authors contributed to the article and approved the submitted version.
Conflict of Interest
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Publisher’s Note
All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.
References
Allen, J. W. (2012). A chemical referral and reference guide to the known species of psilocin and/or psilocybin-containing mushrooms and their published analysis and bluing reactions Ethnomycol. J. 9, 130–175.
Amsterdam, J., van Opperhuizen, A., Brink, W., and van den. (2011). Harmful potential of magic mushroom use: a review. Regul. Toxicol. Pharmacol. 59, 423–429. doi: 10.1016/j.yrtph.2011.01.006
Andersson, C., Kristinsson, J., and Gry, J. (2009). Occurrence and Use of Hallucinogenic Mushrooms Containing Psilocybin Alkaloids. Denmark: Nordic Council of Ministers.
Araujo, R., and Sampaio-Maia, B. (2018). Fungal Genomes and Genotyping,” in Advances in Applied Microbiology. Cambridge: Academic Press Inc, 37–81. doi: 10.1016/bs.aambs.2017.10.003
Arora, D. (1986). Mushrooms demystified : a comprehensive guide to the fleshy fungi. Berkeley: Ten Speed Press.
Badham, E. R. (1984). Ethnobotany of psilocybin mushrooms, especially Psilocybe cubensis. J. Ethnopharmacol. 10, 249–254. doi: 10.1016/0378-8741(84)90007-2
Badotti, F., De Oliveira, F. S., Garcia, C. F., Vaz, A. B. M., Fonseca, P. L. C., Nahum, L. A., et al. (2017). Effectiveness of ITS and sub-regions as DNA barcode markers for the identification of Basidiomycota (Fungi). BMC Microbiol. 17:42. doi: 10.1186/s12866-017-0958-x
Bashir, H., Jabeen, S., Bashir, H., and Khalid, A. N. (2020). Gymnopilus dunensis, a new species from Punjab province, Pakistan. Phytotaxa 428, 51–59. doi: 10.11646/phytotaxa.428.1.5
Borovička, J. (2008). The wood-rotting bluing Psilocybe species in Central Europe - an identification key. Czech Mycol. 60, 173–192. doi: 10.33585/cmy.60202
Borovička, J., Oborník, M., Stříbrný, J., Noordeloos, M. E., Parra Sánchez, L. A., and Gryndler, M. (2015). Phylogenetic and chemical studies in the potential psychotropic species complex of Psilocybe atrobrunnea with taxonomic and nomenclatural notes. Persoonia Mol. Phylogeny Evol. Fungi 34, 1–9. doi: 10.3767/003158515X685283
Boy, A., Lopez, V., Dave, J., Aquino, C., Undan, J. Q., Grace, K., et al. (2016). Molecular identification and phylogeny of some wild microscopic fungi from selected areas of Jaen, Nueva Ecija, Philippines. Adv. Environ. Biol. 10, 153–158.
Carhart-Harris, R. L., Bolstridge, M., Rucker, J., Day, C. M. J., Erritzoe, D., Kaelen, M., et al. (2016). Psilocybin with psychological support for treatment-resistant depression: an open-label feasibility study. Lancet Psychiatry 3, 619–627. doi: 10.1016/S2215-0366(16)30065-7
Castellanos, J. P., Woolley, C., Bruno, K. A., Zeidan, F., Halberstadt, A., and Furnish, T. (2020). Chronic pain and psychedelics: a review and proposed mechanism of action. Reg. Anesth. Pain Med. 45, 486–494. doi: 10.1136/rapm-2020-101273
Chen, Z., Zhang, P., and Zhang, Z. (2014). Investigation and analysis of 102 mushroom poisoning cases in Southern China from 1994 to 2012. Fungal Divers 64, 123–131. doi: 10.1007/s13225-013-0260-7
Cho, S. E., Jo, J. W., Kwag, Y.-N., Lee, H., Chung, J.-W., Oh, S. H., et al. (2021). Complete mitochondrial genome sequence of Gymnopilus junonius. Mitochond. DNA Part B 6, 1020–1021. doi: 10.1080/23802359.2021.1895692
Cortés-Pérez, A., Ramírez-Guillén, F., Guzmán, G., Guzmán-Dávalos, L., Rockefeller, A., and Ramírez-Cruz, V. (2021). Type studies in five species of Psilocybe (Agaricales, basidiomycota). Nov. Hedwigia 112, 197–221. doi: 10.1127/nova-hedwigia/2020/0609
Dalefield, R. (2017). Mycotoxins and Mushrooms. Vet. Toxicol. Aust. NZ 373–419. doi: 10.1016/b978-0-12-420227-6.00020-7
Desjardin, D. E., and Perry, B. A. (2018). The genus Pluteus (Basidiomycota, Agaricales, Pluteaceae) from Republic of São Tomé and Príncipe, West Africa. Mycosphere 9, 598–617. doi: 10.5943/mycosphere/9/3/10
Dos Santos, R. G., Bouso, J. C., and Hallak, J. E. C. (2019). Serotonergic hallucinogens/psychedelics could be promising treatments for depressive and anxiety disorders in end-stage cancer. BMC Psychiatry 19:1–13. doi: 10.1186/s12888-019-2288-z
Duneman, N. (2021). Legalizing Marijuana, Psilocybin Mushrooms, and MDMA for Medical Use. Thesis. Montreal: Concordia University.
Ediriweera, S., Wijesundera, R. L. C., Nanayakkara, C., and Weerasena, J. (2015). First Report of Panaeolus sphinctrinus and Panaeolus foenisecii (Psathyrellaceae, Agaricales) on Elephant Dung from Sri Lanka. Front. Environ. Microbiol. 1:19–23. doi: 10.11648/j.fem.20150102.12
Estrada, E. M., Suárez, M. E., and Maillard, O. (2020). Checklist of Bolivian Agaricales. 1: Species with dark and pink spore prints. Mycotaxon 134, 739–761. doi: 10.5248/134.739
Flaherty, G. T., Maxemous, K. K., Nossier, R. E., and Bui, Y. G. (2017). The highs and lows of drug tourism: A travel medicine perspective. J. Travel Med. 24, 1–3. doi: 10.1093/jtm/tax068
Gartz, J. (1987). Occurrence of psilocybin and baeocystin in fruit bodies of Pluteus salicinus. Planta Med. 53, 290–291. doi: 10.1055/S-2006-962710
Gartz, J. (1996). Magic mushrooms around the world: a scientific journey across cultures and time - the case for challenging research and value systems. New York, NY: LIS Publications.
Gerhardt, E. (1996). Taxonomische Revision der Gattungen Panaeolus und Panaeolina (Fungi, Agaricales, Coprinaceae). Bibl. Bot. 147, 1–149. doi: 10.1007/978-3-319-23534-9_1
Goldman, G. B., and Gryzenhout, M. (2019). Field guide to mushrooms and other fungi of South Africa. 1st ed. New York, NY: Penguin Random House.
Guzmán, G. (1978). Index of taxa in the genus Psilocybe. Taxonomic studies on dark-spored agarics. Mycotaxon 6, 464–476.
Guzmán, G. (1983). The genus Psilocybe. A systematic revision of the known species including the history, distribution and chemistry of the hallucinogenic species. Beihefte zur Nov. Hedwigia. 74, 1–439.
Guzmán, G. (2008). Hallucinogenic mushrooms in Mexico: an overview. Econ. Bot. 62, 404–412. doi: 10.1007/s12231-008-9033-8
Guzmán, G. (2009). “The hallucinogenic Mushrooms: Diversity, Traditions, Use and Abuse with Special Reference to the Genus Psilocybe,” in Fungi from Different Environments, eds J. Misra and S. Deshmukh (New York, NY: Taylor & Francis), 256–276. doi: 10.1201/9780429061653-11
Guzmán, G. (2012). New taxonomical and ethnomycological observations on Psilocybe s.s. (Fungi, Basidiomycota, Agaricomycetidae, Agaricales, Strophariaceae) from Mexico, Africa and Spain. Acta Botaìnica Mex. 100, 79–106. doi: 10.21829/abm100.2012.32
Guzmán, G., Allen, J. W., and Gartz, J. (1998). A worldwide geographical distribution of the Neurotropic fungi, an analysis and discussion. Africa 14, 1–107.
Guzmán, G., Guillén, F. R., Hyde, K. D., and Karunarathna, S. C. (2012). Psilocybe s.s. in Thailand: Four new species and a review of previously recorded species. Mycotaxon 119, 65–81. doi: 10.5248/119.65
Guzmán, G., Kroeger, P., and Ramirez-Guillen, F. (2016). Psilocybe (Basidiomycotina, Agaricales, Strophariaceae) in Canada, with a special review of species from British Columbia. Mycotaxon 3, 179–193.
Guzmán, G., Ramírez, F., Escalona, F., and Jacobs, J. (2004). New hallucinogenic mushrooms in Mexico belonging to the genus Psilocybe (Basidiomycotina, Agaricales, Strophariaceae). Artic. Int. J. Med. Mushrooms 2004:70. doi: 10.1615/IntJMedMushr.v6.i3.70
Guzmán-Dávalos, L., and Herrera, M. (2006). A new bluing, probably hallucinogenic species of Gymnopilus P. Karst. (Agaricomycetideae) from Mexico. Int. J. Med. Mushrooms 8, 289–293. doi: 10.1615/IntJMedMushr.v8.i3.110
Guzmán-Dávalos, L., Mueller, G. M., Cifuentes, J., Miller, A. N., and Santerre, A. (2003). Traditional infrageneric classification of Gymnopilus is not supported by ribosomal DNA Sequence Data. Mycologia 95, 1204–1214. doi: 10.2307/3761920
Halling, R. E., Ammirati, J. F., Traquair, J. A., and Horgen, P. A. (1987). Poisonous Mushrooms of the Northern United States and Canada. Brittonia 39:25. doi: 10.2307/2806968
Hanks, J. B., and González-Maeso, J. (2016). “Hallucinogens: circuits, behavior, and translational models,” in Neuropathology of Drug Addictions and Substance Misuse. Amsterdam: Elsevier Inc, 813–820. doi: 10.1016/B978-0-12-800212-4.00076-5
Hartman, S. (2018). Psilocybin could be legal for therapy by 2021. Available online at: https://www.rollingstone.com/culture/culture-news/psilocybin-legal-therapy-mdma-753946/ [Accessed November 17, 2020].
Hatfield, G., and Valdes, L. (1978). The occurrence of psilocybin in Gymnopilus species. Lloydia 41, 140–144.
He, M. Q., Zhao, R. L., Hyde, K. D., Begerow, D., Kemler, M., Yurkov, A., et al. (2019). Notes, outline and divergence times of Basidiomycota. Fungal Divers 99, 105–367. doi: 10.1007/s13225-019-00435-4
Holec, J. (2005). The genus Gymnopilus (Fungi, Agaricales) in the Czech Republic with respect to collections from other European countries. Acta Musei Natl. Pragae, Ser. B Hist. Nat. 61, 1–52. doi: 10.1127/0029-5035/2008/0087-0001
Holec, J., Antonin, V., Graca, M., and Moreau, P.-A. (2003). Gymnopilus igniculus-find from the Czech Republic and notes on its variability. Czech Mycol. 55, 3–4.
Hopple, J. S., and Vilgalys, R. (1999). Phylogenetic relationships in the mushroom genus Coprinus and dark-spored allies based on sequence data from the nuclear gene coding for the large ribosomal subunit RNA: divergent domains, outgroups, and monophyly. Mol. Phylogenet. Evol. 13, 1–19. doi: 10.1006/mpev.1999.0634
Hosen, M. I., Liang, X., Xu, J., and Li, T. H. (2019). Pluteus squarrosus sp. nov. (Pluteus sect. Celluloderma, Pluteaceae) from northeast China. Nord. J. Bot. 37, 1–7. doi: 10.1111/NJB.02427
Hu, Y., Mortimer, P., Karunarathna, S., Raspé, O., Promputtha, I., Yan, K., et al. (2020). A new species of Panaeolus (Agaricales, Basidiomycota) from Yunnan, Southwest China. Phytotaxa 434, 22–34. doi: 10.11646/phytotaxa.434.1.3
Iliffe, R. (2010). Getting to grips with Pluteus. F. Mycol. 11, 78–92. doi: 10.1016/j.fldmyc.2010.07.005
Ishaq, M., Karunarathna, S. C., Dauner, L., Mai Sci, C. J., Tibpromma, S., Asad, S., et al. (2021). Pluteus cervinus and Laccaria moshuijun (Agaricales, Basidiomycota), New Records from Pakistan. Chiang Mai J. Sci. 48, 909–921.
Johnson, M. W., Griffiths, R. R., Hendricks, P. S., and Henningfield, J. E. (2018). The abuse potential of medical psilocybin according to the 8 factors of the Controlled Substances Act. Neuropharmacology 142, 143–166. doi: 10.1016/j.neuropharm.2018.05.012
Justo, A., Malysheva, E., Bulyonkova, T., Vellinga, E. C., Cobian, G., Nguyen, N., et al. (2014). Molecular phylogeny and phylogeography of Holarctic species of Pluteus section Pluteus (Agaricales: Pluteaceae), with description of twelve new species. Phytotaxa 180, 1–85. doi: 10.11646/phytotaxa.180.1.1
Justo, A., Minnis, A. M., Ghignone, S., Menolli, N., Capelari, M., Rodríguez, O., et al. (2011a). Species recognition in Pluteus and Volvopluteus (Pluteaceae, Agaricales): morphology, geography and phylogeny. 10, 453–479. doi: 10.1007/s11557-010-0716-z
Justo, A., Vizzini, A., Minnis, A. M., Menolli, N., Capelari, M., Rodríguez, O., et al. (2011b). Phylogeny of the Pluteaceae (Agaricales, Basidiomycota): Taxonomy and character evolution. Fungal Biol. 115, 1–20. doi: 10.1016/j.funbio.2010.09.012
Kalichman, J., Kirk, P. M., and Matheny, P. B. (2020). A compendium of generic names of agarics and Agaricales. Taxon 69, 425–447. doi: 10.1002/tax.12240
Karsten, P. A. (1879). Rysslands, Finlands och den Skandinaviska halföns Hattsvampar. Förra Delen: Skifsvampar. Bidr. till Kännedom av Finlands Natur och Folk 32, 1–571.
Kaur, A., Atri, N. S., and Kaur, M. (2014). Diversity of coprophilous species of Panaeolus (Psathyrellaceae, agaricales) from Punjab. India Biodiver. 15, 115–130. doi: 10.13057/biodiv/d150202
Khan, J., Kiran, M., Jabeen, S., Sher, H., and Khalid, A. N. (2017). Gymnopilus penetrans and G. swaticus sp. nov. (Agaricomycota: Hymenogastraceae); a new record and a new species from northwest Pakistan. Phytotaxa 312, 60–70. doi: 10.11646/PHYTOTAXA.312.1.4
Kinge, T. R., Goldman, G., Jacobs, A., Ndiritu, G. G., and Gryzenhout, M. (2020). A first checklist of macrofungi for South Africa. MycoKeys 63, 1–48. doi: 10.3897/mycokeys.63.36566
Kirk, P., Cannon, P., Minter, D., and Stalpers, J. (2008). Dictionary of the Fungi. 10th ed. Wallingford: CAB International.
Krebs, T. S., and Johansen, P. O. (2013). Psychedelics and mental health: a population study. PLoS One 8:e63972. doi: 10.1371/journal.pone.0063972
Landry, B. (2016). Phylogenetic relationships of alpha-amanitin producing Galerina from British Columbia. Thesis. Vancouver: University of British Columbia, doi: 10.14288/1.0378696
Lee, S., Ryoo, R., Choi, J. H., Kim, J. H., Kim, S. H., and Kim, K. H. (2020). Trichothecene and tremulane sesquiterpenes from a hallucinogenic mushroom Gymnopilus junonius and their cytotoxicity. Arch. Pharm. Res. 43, 214–223. doi: 10.1007/S12272-020-01213-6
Lenz, C., Wick, J., Braga, D., GarciaAltares, M., Lackner, G., Hertweck, C., et al. (2020). Injury-Triggered Blueing Reactions of Psilocybe “Magic” Mushrooms. Angew. Chemie Int. 59, 1450–1454. doi: 10.1002/anie.201910175
Li, S., Ma, Q.-B., Tian, C., Ge, H.-X., Liang, Y., Guo, Z.-G., et al. (2019). Cardiac arrhythmias and cardiac arrest related to mushroom poisoning: A case report. World J. Clin. Cases 7, 2330. doi: 10.12998/WJCC.V7.I16.2330
Li, Y.-K., Yuan, Y., and Liang, J.-F. (2014). Morphological and molecular evidence for a new species of Psilocybe from southern China. Mycotaxon 129, 215–222. doi: 10.5248/129.215
Liu, M., and Bau, T. (2019). Gymnopilus minisporus sp. nov., a new species and a new record of the European species G. hybridus from northeast China. Phytotaxa 397, 159–168. doi: 10.11646/PHYTOTAXA.397.2.3
Ma, T. (2014). Taxonomy of Psilocybe s.l. and Panaeolus in Yunnan, Southwest China, with notes on related genus Protostropharia. Beijing: Chinese Academy of Forestry.
Ma, T., Feng, Y., Lin, X. F., Karunarathna, S. C., Ding, W. F., and Hyde, K. D. (2014). Psilocybe chuxiongensis, a new bluing species from subtropical China. Phytotaxa 156, 211–220. doi: 10.11646/phytotaxa.156.4.3
Ma, T., Ling, X. F., and Hyde, K. D. (2016). Species of Psilocybe (Hymenogastraceae) from Yunnan, Southwest China. Phytotaxa 284, 181–193. doi: 10.11646/phytotaxa.284.3.3
Maeta, K., Ochi, T., Tokimoto, K., Shimomura, N., Maekawa, N., Kawaguchi, N., et al. (2008). Rapid species identification of cooked poisonous mushrooms by using real-time PCR. Appl. Environ. Microbiol. 74, 3306–3309. doi: 10.1128/AEM.02082-07
Malysheva, E., Moreno, G., Villarreal, M., Malysheva, V., and Svetasheva, T. (2019). The secotioid genus Galeropsis (Agaricomycetes, Basidiomycota): a real taxonomic unit or ecological phenomenon? Mycol. Prog. 18, 805–831. doi: 10.1007/s11557-019-01490-6
Marlan, D. (2019). Beyond cannabis: psychedelic decriminalization and social justice. Lewis Clark Law Rev. 23, 851–854.
Maruyama, T., Kawahara, N., Yokoyama, K., Makino, Y., Fukiharu, T., and Goda, Y. (2006). Phylogenetic relationship of psychoactive fungi based on rRNA gene for a large subunit and their identification using the TaqMan assay (II). Forensic Sci. Int. 163, 51–58. doi: 10.1016/j.forsciint.2004.10.028
Maruyama, T., Shirota, O., Kawahara, N., Yokoyama, K., Makino, Y., and Goda, Y. (2003). Discrimination of psychoactive fungi (commonly called “magic mushrooms”) based on the DNA sequence of the internal transcribed spacer region. J. Food Hyg. Soc. Japan 44, 44–48. doi: 10.3358/shokueishi.44.44
Matheny, P. B., Curtis, J. M., Hofstetter, V., Aime, M. C., Moncalvo, J.-M., Ge, Z.-W., et al. (2006). Major clades of Agaricales: a multilocus phylogenetic overview. Mycologia 98, 982–995. doi: 10.1080/15572536.2006.11832627
McKernan, K., Kane, L. T., Crawford, S., Chin, C.-S., Trippe, A., and McLaughlin, S. (2021). A draft sequence reference of the Psilocybe cubensis genome. F1000 Res. 10:281. doi: 10.12688/f1000research.51613.1
Menolli, N., Asai, T., Capelari, M., and NelsonMenolli, M. (2010). Records and new species of Pluteus from Brazil based on morphological and molecular data. Mycology 1, 130–153. doi: 10.1080/21501203.2010.493531
Menolli, N., Justo, A., and Capelari, M. (2015). Phylogeny of Pluteus section Celluloderma including eight new species from Brazil. Mycologia 107, 1205–1220. doi: 10.3852/14-312
Menolli, N., Justo, A., Arrillaga, P., Pradeep, C. K., Minnis, A. M., and Capelari, M. (2014). Taxonomy and phylogeny of Pluteus glaucotinctus sensu lato (Agaricales, Basidiomycota), a multicontinental species complex. Phytotaxa 188, 78–90. doi: 10.11646/phytotaxa.188.2.2
Meyer, W., Irinyi, L., Hoang, M. T. V., Robert, V., Garcia-Hermoso, D., Desnos-Ollivier, M., et al. (2019). Database establishment for the secondary fungal DNA barcode translational elongation factor 1α (TEF1α). Genome 62, 160–169. doi: 10.1139/gen-2018-0083
Minnis, A. M. (2008). A Systematic Study Of The Euagaric Genus Pluteus Emphasizing Section Celluloderma In The U.S.A. 1st ed. Seattle: Pacific Northwest Fungi Project.
Minnis, A. M., Sundberg, W. J., Methven, A. S., Sipes, S. D., and Nickrent, D. L. (2006). Annulate Pluteus species, a study of the genus Chamaeota in the United States. Mycotaxon 96, 31–39.
Moncalvo, J.-M., Vilgalys, R., Redhead, S. A., Johnson, J. E., James, T. Y., Aime, M. C., et al. (2002). One hundred and seventeen clades of euagarics. Mol. Phylogenet. Evol. 23, 357–400. doi: 10.1016/S1055-7903(02)00027-1
Mullineux, T., and Hausner, G. (2009). Evolution of rDNA ITS1 and ITS2 sequences and RNA secondary structures within members of the fungal genera Grosmannia and Leptographium. Fungal Genet. Biol. 46, 855–867. doi: 10.1016/j.fgb.2009.08.001
Mushroom Observer (2021). Available online at: https://mushroomobserver.org/ [Accessed October 7, 2021].
Musshoff, F., Madea, B., and Beike, J. (2000). Hallucinogenic mushrooms on the German market - Simple instructions for examination and identification. Forensic Sci. Int. 113, 389–395. doi: 10.1016/S0379-0738(00)00211-5
NCBI (2021). The NCBI Taxonomy database (Nucleotide). Available online at: https://www.ncbi.nlm.nih.gov/nuccore [Accessed October 7, 2021].
Norvell, L. L., Hawksworth, D. L., Petersen, R. H., and Redhead, S. A. (2010). IMC9 Edinburgh Nomenclature Sessions. IMA Fungus 1, 143–147. doi: 10.5598/imafungus.2010.01.02.05
O’Hanlon, R. (2018). “Fungi in the Environment,” in Fungi: Biology and ApplicationsThird Edition, ed. K. Kavanagh New York, NY: Wiley-Blackwell. 333–355. doi: 10.1002/9781119374312
Ola’h, G. M. (1969). Le genre Panaeolus: Essai taxinomique et physiologique. Louis Marie: Paris Herb, 116–119.
Osmundson, T. W., Robert, V. A., Schoch, C. L., Baker, L. J., Smith, A., Robich, G., et al. (2013). Filling gaps in biodiversity knowledge for macrofungi: contributions and assessment of an herbarium collection DNA Barcode Sequencing Project. PLoS One 8:62419. doi: 10.1371/journal.pone.0062419
Oss, O. T., and Oeric, O. N. (1991). “Psilocybin : magic mushroom grower’s guide,” in A handbook for psilocybin enthusiasts (San Francisco: Quick American Pub), 1–81. doi: 10.1520/jfs2004276
Oxford Analytica (2021). Psychedelic drug research will increase globally. Expert Brief. 2021:9. doi: 10.1108/OXAN-DB260709
Passie, T., Seifert, J., Schneider, U., and Emrich, H. M. (2002). The pharmacology of psilocybin. Addict. Biol. 7, 357–364. doi: 10.1080/1355621021000005937
Peintner, U., Bougher, N. L., Castellano, M. A., Moncalvo, J. M., Moser, M. M., Trappe, J. M., et al. (2001). Multiple origins of sequestrate fungi related to Cortinarius (Cortinariaceae). Am. J. Bot. 88, 2168–2179. doi: 10.2307/3558378
Quélet, L. (1872). Les Champignons du Jura et des Vosges. Mémoires la Société d’Émulation Montbéliard 2. 5, 43–332.
Ramírez-Cruz, V., Guzmán, G., and Guzmán-Dávalos, L. (2012). New Combinations in the genus Deconica (Fungi, Basidiomycota, Agaricales). Sydowia 64, 217–219.
Ramírez-Cruz, V., Guzmán, G., and Guzmán-Dávalos, L. (2013a). Type studies of Psilocybe sensu lato (Strophariaceae, Agaricales). Sydowia 2, 277–319.
Ramírez-Cruz, V., Guzmán, G., Villalobos-Arámbula, A. R., Rodríguez, A., Matheny, P. B., Sánchez-García, M., et al. (2013b). Phylogenetic inference and trait evolution of the psychedelic mushroom genus Psilocybe sensu lato (Agaricales). Botany 91, 573–591. doi: 10.1139/cjb-2013-0070
Razaq, A., Khalid, A. N., and Illyas, S. (2012). Molecular identification of Lyophyllum connatum and Paneolus shinctrinus (Basidiomycota, Agaricales) from Himalyan moist temperature forests of Pakistan. Int. Agricault. Biol. 14, 1001–1004.
Redhead, S. A., and Guzmán, G. (1985). The genus Psilocybe. a systematic revision of the known species including the history, distribution and chemistry of the hallucinogenic species. Mycologia 77:172. doi: 10.2307/3793267
Redhead, S. A., Moncalvo, J.-M., Vilgalys, R., Matheny, P. B., and Guzmán, L. Guzmán-Dávalos. (2007). Proposal to Conserve the Name Psilocybe (Basidiomycota) with a Conserved Type. Int. Assoc. Plant Taxon. 56, 255–257. doi: 10.2307/25065762
Rees, B. J., Marchant, A., and Zuccarello, G. C. (2004). A tale of two species—possible origins of red to purple-coloured Gymnopilus species in Europe. Australas. Mycol. 22, 57–72.
Rees, B. J., Orlovich, D. A., and Marks, P. B. D. (1999). Treading the fine line between small-statured Gymnopilus and excentrically stipitate Galerina species in Australia. Mycol. Res. 103, 427–442. doi: 10.1017/S095375629800745X
Reiff, C. M., Richman, E. E., Nemeroff, C. B., Carpenter, L. L., Widge, A. S., Rodriguez, C. I., et al. (2020). Psychedelics and psychedelic-assisted psychotherapy. Am. J. Psychiatry 177, 391–410. doi: 10.1176/appi.ajp.2019.19010035
Reingardiene, D., Vilcinskaite, J., and Lazauskas, R. (2005). Hallucinogenic mushrooms. Medicina 41, 1067–1070.
Rodriguez, O., Galva-Corona, A., Villalobos-Arambula, A., Vargas, G., and Guzmán-Dávalos, L. (2008). Pluteus horakianus, a new species from Mexico, based on morphological and molecular data. Sydowia 61, 39–52.
Rogan, J. (2018). Joe Rogan Experience Podcast #1169 with Elon Musk. Available online at: https://sonix.ai/r/gf7b9enjv94nkrpJ1KqEa8mk/transcript.pdf (accessed May 9, 2022).
Rogan, J. (2019). Joe Rogan Experience Podcast #1306. Available online at: https://www.youtube.com/watch?v=fTAPaHcLAhw (accessed May 31, 2019).
Rumack, B. H., and Spoerke, D. G. (1994). Handbook of Mushroom Poisoning: Diagnosis and Treatment. Boca Raton: CRC Press.
Saupe, S. G. (1981). Occurrence of psilocybin/psilocin in Pluteus salicinus Pluteaceae. Mycologia 73, 781–784. doi: 10.2307/3759505
Schoch, C. L., Seifert, K. A., Huhndorf, S., Robert, V., Spouge, J. L., Levesque, C. A., et al. (2012). Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA barcode marker for Fungi. Proc. Natl. Acad. Sci. USA 109, 6241–6246. doi: 10.1073/pnas.1117018109
Senn-Irlet, B., Nyffenegger, A., and Brenneisen, R. (1999). Panaeolus bisporus - An adventitious fungus in central Europe, rich in psilocin. Mycologist 13, 176–179. doi: 10.1016/S0269-915X(99)80107-4
Sette, L. D., Passarini, M. R. Z., Rodrigues, A., Leal, R. R., Simioni, K. C. M., Nobre, F. S., et al. (2010). Fungal diversity associated with Brazilian energy transmission towers. Fungal Divers 44, 53–63. doi: 10.1007/s13225-010-0048-y
Ševčíková, H., Borovička, J., and Gates, G. (2021). Pluteus hubregtseorum (Pluteaceae), a new species from Australia and New Zealand. Phytotaxa 496, 147–158. doi: 10.11646/phytotaxa.496.2.4
Shaffer, R. L., and Singer, R. (1976). The Agaricales in modern taxonomy. Mycologia 68:447. doi: 10.2307/3759020
Singer, R. (1958). Monographs of South American Basidiomycetes, especially those of the east slope of the Andes and Brazil. I. The genus Pluteus in South America. Lloydia 21, 195–299.
Spring, M. G., Ostrow, R. D., and Hallock, R. M. (2016). “A Profile of Those Who Use Hallucinogenic Mushrooms,” in Neuropathology of Drug Addictions and Substance Misuse (Amsterdam: Elsevier Inc), 794–800. doi: 10.1016/B978-0-12-800212-4.00074-1
Stríbrný, J., Borovicka, J., and Sokol, M. O. (2003). Psilocybin a psilocinu v nĕkterých druzích hub [Levels of psilocybin and psilocin in various types of mushrooms]. Soud Lek. 48, 45–49.
Tekpinar, A. D., and Kalmer, A. (2019). Utility of various molecular markers in fungal identification and phylogeny. Nov. Hedwigia 109, 187–224. doi: 10.1127/nova_hedwigia/2019/0528
Thomas, K., Peintner, U., Moser, M., and Manimohan, P. (2002). Anamika, a new mycorrhizal genus of Cortinariaceae from India and its phylogenetic position based on ITS and LSU sequences. Mycol. Res. 106, 245–251. doi: 10.1017/S0953756201005445
Tóth, A., Hausknecht, A., Krisai-Greilhuber, I., Papp, T., Vágvölgyi, C., and Nagy, L. G. (2013). Iteratively refined guide trees help improving alignment and phylogenetic inference in the mushroom family Bolbitiaceae. PLoS One 8:e56143. doi: 10.1371/journal.pone.0056143
Tyls, F., Palenicek, T., and Horacek, J. (2016). “Neurobiology of the Effects of Psilocybin in Relation to Its Potential Therapeutic Targets,” in Neuropathology of Drug Addictions and Substance Misuse (Amsterdam: Elsevier Inc), 782–793. doi: 10.1016/B978-0-12-800212-4.00073-X
Van der Walt, R., Dames, J., and Hawley-MacMaster, G. (2020). Fungi and Lichens of the Limpopo Valley & Mapungubwe National Park. Limpopo: Retha van der Walt.
Wang, Y.-W., and Tzean, S.-S. (2015). Dung-associated, Potentially Hallucinogenic Mushrooms from Taiwan. Taiwania 60, 160–168. doi: 10.6165/tai.2015.60.160
Watling, R. (1977). A Panaeolus poisoning in scotland. Mycopathologia 61, 187–190. doi: 10.1007/BF00468015
Wesselink, A. (2018). DNA markers for forensic identification of non-human biological traces. Amsterdam: University of Amsterdam.
Wijayawardene, N., Hyde, K., Al-Ani, L. K. T., Tedersoo, L., Haelewaters, D., Rajeshkumar, K. C., et al. (2020). Outline of Fungi and fungus-like taxa. Mycosphere 11, 1060–1456. doi: 10.5943/mycosphere/11/1/8
Wurst, M., Kysilka, R., and Flieger, M. (2002). Psychoactive tryptamines from basidiomycetes. Folia Microbiol 47, 3–27. doi: 10.1007/BF02818560
Keywords: hallucinogenic mushrooms, taxonomy, phylogeny, Psilocybe, Panaeolus, Pluteus, Gymnopilus, psychedelic mushrooms
Citation: Strauss D, Ghosh S, Murray Z and Gryzenhout M (2022) An Overview on the Taxonomy, Phylogenetics and Ecology of the Psychedelic Genera Psilocybe, Panaeolus, Pluteus and Gymnopilus. Front. For. Glob. Change 5:813998. doi: 10.3389/ffgc.2022.813998
Received: 12 November 2021; Accepted: 07 April 2022;
Published: 23 May 2022.
Edited by:
Janette Bulkan, University of British Columbia, CanadaReviewed by:
Braham Dhillon, University of Florida, United StatesPeter Matthew Scott, Department of Primary Industries and Regional Development of Western Australia (DPIRD), Australia
Copyright © 2022 Strauss, Ghosh, Murray and Gryzenhout. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Marieka Gryzenhout, gryzenhoutm@ufs.ac.za