The study was conducted in the communities of San José Buenavista and Agua Ancha, in the municipality of Eloxochitlán de Flores Magón. Both communities are located in the north of the state of Oaxaca, Mexico (Fig. 1). The study area is located between the geographical coordinates 18°09’ and 18°15’N, and 96°50’ and 96°55’W at 1460 m.a.s.l. According to Köppen’s classification, modified by García (2004), there are two predominant climate types in the area: (A)C(m) (semi-warm sub-humid), with average temperatures above 18°C and abundant summer rains, and (A)C(fm) (temperate semi-warm humid) with rains all year, with a temperature of -3°C in cold months and an average temperature above 18°C. Based on Rzedowski’s (2006) classification, the predominant types of vegetation in the municipality are mountain cloud forest and evergreen tropical forest. In the mountain cloud forest, the presence of Liquidambar styraciflua, Pinus teocote, Quercus spp., and species of the families Orchidaceae, Bromeliaceae, Cyatheaceae y Piperaceae was recorded. Meanwhile in the evergreen tropical forest, the vegetation included Brosimum alicastrum, Quercus spp., Licania platypuss, and Manilkara zapota.

A field study was conducted during the rainy seasons from 2017 to 2021. A total of 291 non-structured, semi-structured, and structured interviews were carried by using the “snowball” technique (Sandoval, 2002). The following information was obtained from the conducted interviews: i) socio-demographic data disaggregated by gender, age, language, inhabitants per household, level of schooling, and economic activities; ii) known wild fungi, including information about their usages, local naming, local classification, ecology, morphology and their conceptualization of a fungus; iii) transmission mechanisms of mycological knowledge; iv) methods for mushroom picking; v) traditional culinary use; vi) mushroom poisoning; and vii) relationship among fungi and wild fauna. All mentioned fungi in the interviews were inventoried according to the methods proposed by Ruan et al. (2004), Garibay et al. (2006), Burrola et al. (2012) and Domínguez et al. (2015). The method consisted in recording all the traditional names in Mazatec language of the organisms that the interviewees consider to be mushrooms. Along with the names, drawings and verbal detailed descriptions were requested to the interviewees; and when necessary mushroom photographs were used. Finally, the identity of the species was confirmed with fresh samples collected in the field for all species. In order to select the people to be interviewed, two techniques were used: i) “snowball”, which consists of identifying people with deep traditional ethnomycological knowledge; afterward asking to this people if they knew other people which also had broad mycological knowledge; then we contacted this new people (Sandoval, 2002), and ii) randomly within the study community, knocking from door to door, assuming that knowledge related to the mycological resource in the community was widely distributed. To recognize the taxonomic identity of the fungi mentioned in the interviews, the methodology proposed by Garibay et al. (2006) and Burrola et al. (2012) was used, which consists of using picture sets of fungi from the region and also to gather fresh specimens along with the local expert people from the studied community.

To establish an importance score for the wild mushrooms used by the community inhabitants, we used the Frequency of Mention Index proposed by Garibay et al. (2007) which was directly obtained from the free list of fungi known in each of the interviews. Mention Index (MI) was obtained by dividing the number of mentions of a fungus by the number of informants and multiplying this number by 10. The frequency of mention (FM) was obtained by dividing the number of total mentions of a fungus by the number of informants.

Additionally, fresh specimens were collected with the collaboration and expertise of local inhabitants or were purchased at the local markets of the studied region. Collected mushrooms were photographed and their macro- and micromorphological features were described. Taxonomic identification was performed by using specific literature (Anderson & Ullrich, 1979; Largent et al., 1980; Guzmán, 1983; Torres et al., 2015; Herrera et al., 2018; Fang et al., 2021; Rodríguez et al., 2022). The scientific nomenclature of specimens was based on Index Fungorum (http://www.indexfungorum.org). Dehydrated labelled specimens were deposited in the Mycological Collection of the Microbiology Area of the Postgraduate College (COLPOS) in Montecillo, State of Mexico. Mazatec language writing and its translation was carried out with the support of local translators and specialized dictionaries of the linguistic variant of Eloxochitlán de Flores Magón (Cházarez, 2012; Avendaño & Agee, 2013; Avendaño et al., 2013).

Twenty-seven species of wild fungi with biocultural relevance were collected and identified within the study community (Table 1), of which 18 are edible, five have medicinal usages, and five are used for ludic purposes (one of the species has two different uses). The name to identify fungal species is the word “tjii̱n”, which means “mushroom” (or “thick”) followed by an adjective or noun. Fungal species are classified into three groups according to their attributed use: i) edible species are named tjii̱n ra ma chine, or “mushrooms that can be eaten”, ii) medicinal species are called tjii̱n ra ma xkii̱, which literally means mushrooms that “make you feel good”, ”make you feel warm inside” or “medicinal mushrooms”, and iii) species that do not have any particular relevance are named tjii̱n ra siskajinnáem> or tjii̱n ska, which literally means “mushrooms that drive you crazy” or “crazy fungi”.

The fungal species mentioned by the interviewed people are classified into six ethnotaxons that correspond to: i) white fungi (tjii̱n chroba) including Pleurotus spp. and Hydnopolyporus fimbriatus, ii) ear fungi (tjii̱n chíká) including Auricularia spp.; iii) oak fungi (tjii̱n ítá) including Cantharellus spp. and Laetiporus sulphureus; iv) medicinal mushrooms (tjii̱n xkii̱) including Psilocybe spp. and Panaeolus cyanescens; v) “hard texture fungi” (tjii̱n tája) including Ganoderma spp., H. fimbriatus and Lentinus crinitus; and vi) dust fungi (tjii̱n chijo) including Geastrum lageniforme and Calvatia cyathiformis.

The importance of mushrooms for the community inhabitants lies in their edible value. A significant percentage of people (42%) mentioned that mushrooms are a better food than meat or any other food; some others mentioned that mushrooms have a good taste (25%), others did not provide any answer (23%), few mentioned their medicinal usages (6%) and very few (4%) said that mushrooms were beneficial to the earth and nature, or that they were not relevant. Among community inhabitants, mushrooms are conceptualized differently; 42% consider that mushrooms are plants, even when Mazatec language distinguishes plants from mushrooms and has different words for each of them; 19% ignore what a mushroom is, and did not provide an answer to this question; and 9% define a mushroom as “…something that emerges from the forest tree trunks, being the trunk itself that takes that form…”. Interestingly 26% of interviewed people conceive fungi with some fascinating interpretations for example as: “… the juice or the soul of the trees”, “a juice coming from the earth”, “something that is born from a root or plant”, “an animal”, “a medication”, “a putrid stick”, “dry leaves”, “bugs”, “the hard outer covering of the trees that turns into a mushroom”, “something that came from an animal”, “a decomposer plant (because is something that decay dead trees)”, “something that came from the heart of the trees”, “a pest from the trees that grow in there”, “something that came from mother earth”, “butterflies debris that turns into mushrooms”, “plants which are from the family of lichens”, “a seed”, “manna (an edible food that God sent to the Israelites according to Christian people)”, “soul in purgatory (according to Catholic religion, it refers to the souls of the dead who are in purgatory)” and “a kind of orchid”.

The traditional knowledge used to identify wild edible mushrooms is transmitted by oral tradition from adults to children (Fig. 2c). As well, the exchange of mycological knowledge is a bidirectional process that takes place when a new nuclear family (created when new members are added to a family, either from community members or people from other cultures) is created. Knowledge transfer includes identification of the type of substrate where a specific mushroom grows, vegetation types where some mushrooms can be found, forest mushroom species with high biocultural importance, seasonality, and traditional names. In addition, people use to communicate some procedures to identify if a wild mushroom is edible, such as tasting a small mushroom fragment and if it has an unpleasant or peppery (spicy) taste, then it is not edible. A small proportion of interviewed people (2%) recommend rubbing a piece of garlic on the fungus and if it turns black, it demonstrates that is not an edible mushroom. However, is important to note that this last procedure is not scientifically accurate and might cause either gastrointestinal mycetism or be lethal.

Wild mushrooms are picked up in nearby forests and sometimes also in coffee plantations. When the collection of wild edible mushrooms is abundant, and they are not used for sale, people use to give a small amount of them to their relatives or neighbors as a gift which strengthens the social relationships among the community members. Regarding picking practices, 56% of interviewees use techniques that promote the fructification of sporomes in subsequent years. These techniques consist of picking mature specimens and leaving part of the stipe on the ground, preventing mycelium disturbance. Conversely, a smaller number of people use inadequate harvesting practices that may impact the presence of sporomes in subsequent years. These practices include pulling sporomes out from the stipe base, thus carrying the soil and the mycelium with them. Another relevant aspect that may affect the sporome fructification in subsequent years is their transportation method. People prefer transporting wild mushrooms using broad leaves or traditional hampers called “chitates” (ínchjro). This transport method is an appropriate harvesting practice because it promotes spore dispersion (Fig. 2g). However, wild mushrooms are sometimes also carried out in plastic bags or buckets, that prevent sporome spore dispersal (Fig. 2h).

We were informed of 24 cases of mycetism during the interviews. Three of these cases were lethal. Regarding non-lethal poisoning cases (21), the symptoms were vomiting, stomach pain, rash, hallucinations, and gastroenteritis. Unfortunately, it was not possible to identify the species of fungi responsible for the poisoning because inhabitants only describe and know the fungus colour and, sometimes, the substrate where they grew. The oldest mycetism case among community inhabitants recorded in the interviews dates back to 1970. The interview said, “My granny loved to eat wild mushrooms! One day, she went out looking… and saw some that looked tasty to eat. My mom told her that those mushrooms might hurt her because they did not look good to be eaten. My granny ignored her and prepared them as Tjii̱n chroba. She ate all of them herself. In the late afternoon, she was complaining of stomach pain and vomiting, but she recovered the next day. I think she removed all the bad stuff when she vomited. My mom scolded her and told her not to eat wild mushrooms carelessly”. The most recent mycetism case occurred in 2016 when a person, following the advice of someone, collected white wild mushrooms and ate them with the family. As a consequence of the poisoning, the family restricted their wild mushroom consumption to Cantharellus spp. and Pleurotus sp. Nowadays, the parent is the only person who consumes Pleurotus sp. after a detailed inspection of the sporomes.

According to the MI, the most important mushroom species for the inhabitants are Pleurotus spp. (MI=9.73) Auricularia spp. (MI=9.42), S. commune (MI=8.66) and S. radiatum (MI=8.66). The species with the lower MI correspond to P. albidus (MI=0.65) and C. aff. confluens (MI=0.62). Meanwhile, the most mentioned species for ludic use was A. nigricans (MI=2.03), whereas G. lageniforme obtained the lowest mentions (MI=0.10). P. cubensis (MI=8.11) was the most important species for medicinal purposes, because it is frequently used in ceremonies whose purpose is to heal people. P. cyanescens and P. yungensis (MI=0.03) showed the lowest MI. It is considered that P. cyanescens use is rare because it was only mentioned during one field trip accompanied by an “honguera” (name given in Spanish to women who gather wild mushrooms). P. yungensis is currently used only scarcely and one of the reasons is because it is considered necessary to use many sporomes to have an entheogenic effect. Additionally, this species is included in a single ethnotaxa together with P. mexicana, hindering the differentiation among them.

There is a wide variety of culinary uses for wild mushrooms in the community of study. The dishes and their preparation are:

• “Tesmole” a typical dish of the region (Fig. 3a), which is prepared using nixtamalized dough, achiote seeds (Bixa orellana L.), garlic (Allium sativum L.), onion (A. cepa L.), tomato (Solanum lycopersicum L.), and chilli (Capsicum spp.). Some people prefer not to use garlic and onion because they believe that mushrooms lose their flavor when these ingredients are added. Wild mushrooms used to prepare tesmole are Pleurotus spp. and Cantharellus spp.;
• Roasted mushrooms with “hoja santa” (Piper auritum K.) are prepared when the amount of collected mushrooms is limited. This dish is prepared by placing a base of “platanillo” leaves (Heliconia sp.) and then a base of “hoja santa” leaves. Subsequently, mushrooms, sliced onion, chilli and salt are added. Finally, all the ingredients are sprinkled with water, wrapped using the leaves and placed directly on the coals or on the griddle (Fig. 3b);
• Roasted mushrooms with salt (Fig. 3c) are prepared using S. commune. The mushrooms are soaked in salted water for 15 min and dripped out. Finally, they are roasted directly on the griddle;
• Mexican-style mushrooms (Fig. 3d) are prepared using garlic, onion, tomato and chilli. The mushrooms more frequently used to prepare this dish are Pleurotus spp. and the three species of Cantharellus;
• Mushroom “tacos” are prepared by placing cooked mushrooms (“tesmole”, roasted mushrooms or Mexican-style mushrooms) on a “tortilla” made from corn;
• Roasted mushrooms sprinkled with some salt (Fig. 3e). Collected mushrooms are cooked directly on the griddle and seasoned with salt. The species commonly used to prepare this dish are Daldinia sp., L. crinitus, and Schizophyllum spp., and
• Mushroom “tamales” are cooked using nixtamalized dough mixed with pig lard, and stuffed with “tesmole”, Mexican-style mushrooms or U. maydis. If U. maydis is used, it must be previously cooked with garlic, onion and chilli. All the ingredients are wrapped using corn leaves or platanillo leaves and steamed in a pot.
• According to the information provided by the interviewers, Daldinia sp., A. tremellosa and A. nigricans are species that can be eaten raw. Pleurotus spp. and A. tremellosa can be consumed if they are previously tanned with salt. The preparation consists in wrapping the mushrooms using “hoja santa”. Although A. nigricans is an edible species, only a small group of people consume it. Many interviewed people recognized it but did not consume it. Some others consider it non-edible.

Ludic uses of wild mushrooms are diverse and not restricted to children, adults also play with mushrooms. Children used to play with A. nigricans by placing them behind their ears to make them resemble those of an animal. Children and adults used to play with G. lageniforme and C. cyathiformis sporomes. The mushroom G. lageniforme is squeezed using the fingers or a small stick to see the spores come out. Ludic uses of C. cyathiformis depend on its maturity stages. Mature specimens are kicked or thrown to see the spores come out. As immature specimens resemble a cake, children use the mushroom to play as if they were cookers and they are slicing a cake. G. zonatum is a fungus that can be easily recognized in the forest due to its large size and abundance. It was mentioned during the interviews on several occasions but it was interesting that one person mentioned its use as an ornamental species in the garden.

The species used for medicinal purposes are P. cubensis, P. caerulescens, P. cyanescens, P. mexicana and P. yungensis. Among medicinal mushrooms, interviewees mentioned that small amounts of P. caerulescens produce very strong effects, in terms of hallucinations or “dreams”. These mushrooms are consumed in a ceremony where some rituals are performed. These rituals are a syncretism between pre-Hispanic and Catholic cosmogonies. Before and after the ceremony, it is necessary to have certain kinds of self-care or to follow some specific recommendations for all family members. The ceremony is always performed at night. People who consume mushrooms consider them sacred and have a deep feeling of great respect and love for them. Sacred mushrooms are used for healing, to seek an answer to a better destiny, and to heal spiritually, emotionally or mentally. Medicinal mushrooms are also used to identify people who, through a strong feeling of envy toward another person, harm someone (cursing or wishing the worst to a person). The interviewed people mentioned that when they are having harsh times in their lives, they go with a local chaman in order to know if somebody is responsible for this. Then, they participate in a sacred ceremony where they consume the medicinal mushrooms and they experience clear visions where they are able to literally see the people who is trying to damage them. Some people even mentioned that they have to have an “spiritual fight” with this people during the ceremony in order to stop originating them some damage in their lives. Recreational uses for the genus Psilocybe were not reported by the interviewed people, who by the opposite expressed a deep respect for the sacred mushrooms.

As an element of Mazatec cosmovision, there are customs and beliefs related to wild mushrooms. These beliefs associate various aspects of people’s lives and the surrounding environment to wild mushrooms. Some beliefs relate mushroom abundance to different events. A fructification and abundance indicator for Pleurotus species is the moon. People think when the moon is visible during the day or when there is a crescent moon or full moon, the genus Pleurotus will fruit more abundantly (Fig. 2e). Events such as lightning in a rain or storm are associated with the fructification of the genera Pleurotus and Psilocybe. Regarding to climatic conditions, L. sulphureus is used as an indicator (bioindicator) of a colder winter. Some people believe that their presence or actions may influence the growth of wild mushrooms. When someone finds an immature edible mushroom, they usually pull their earlobe. It is believed that this action will allow the fungus to grow and reach its maturity stage; otherwise, the fungus will pass directly into a decomposing stage without reaching its maturity. Some interviewees mentioned that having sexual intercourse the day they consumed wild mushrooms or cooking the mushrooms with oil will affect the fructification or abundance of wild mushrooms in future times, especially for Pleurotus spp. Other beliefs are based on the idea that wild mushrooms may exert an influence on people’s actions when consumed. There are some beliefs regarding Daldinia sp. (Fig. 2a), it is believed that people who consume large amounts of this fungus may lose their teeth. Some others believe that children born from women who consume large amounts of Daldinia sp. will cry a lot during their childhood. The crying of children is attributed to the noise produced by these mushrooms when cooking. However, people mentioned that crying children can be calmed by feeding them the same fungus. Finally, people believe that the possibility of encountering snakes (Suborder Serpents) instead of wild mushrooms is increased if a person exclusively decides to gather mushrooms or if a person points out that the taste of mushrooms is like eating meat. Specific phrases referring to the fungus as meat such as “this meat is delicious!” or “today we are going to eat meat” are considered an important factor that increases the finding of snakes. Additionally, some persons mentioned that C. cibarius s.l. is guarded by snakes. Some inhabitants used to tell a short legend about C. fusiformis, which is a non-edible fungus. This fungus is known as jno̱ tsálá or jno̱ nisen in the Mazatec language which means lizard “milpa” or mouse “milpa” (Fig. 2f). The legend mentions that lizards (suborder Lacertilia) or mice (Mus spp.) come out of their burrow to till the land and cultivate its own “milpa”. In this case, the crop refers to the mushroom. The mice use to have their milpas on the ground and lizard on the stones. This legend, and beliefs, show the complex links and relevance of mushrooms in the cosmogony of Mazatec culture consequence of centuries of interactions between fungi and people.

Forty-one per cent of interviewed people mentioned that they know that animals eat mushrooms. Mentions include mammals, insects, insect larvae, birds and reptiles. A total of 18 different species were identified. According to the (MI), the animals that use to eat mushrooms are larvae of different animals, colloquially known as “worms” (MI=1.55), reptiles of the Suborder Serpents (MI==1.17), birds (MI=0.65), armadillos (MI=3.34), fruit fly (MI=0.34), squirrels (MI=0.27), rodents (Cricetidae MI=0.21), tepezcuintle (Cuniculus paca, MI=0.17), Tlacuache (opossum, Didelphis sp., MI=0.14), slugs (Trigonochlamydidae, MI=0.07), raccoons (MI=0.07), skunks (Conepatus sp., MI=0.07), badgers (Meles spp., MI=0.07), and ants (MI=0.07). The animals with the lowest MI were goats (MI=0.03) and brocket deers (Mazama temama, MI=0.03). Some inhabitants use rotted fungus to feed their hens or sometimes they use cooked mushrooms to feed their dogs. A person mentioned that his cat used to eat repeatedly certain mushrooms that caused drunkenness.

Wild mushrooms are commercialized from house to house and in the local markets. Among interviewed people, 19.5% dedicate to commercializing wild mushrooms. The species with higher commercial demand is Cantharellus spp., but the most traded and abundant species belong to the Pleurotus genus. A. mellea is the least traded fungus. The mushrooms are sold in containers, used as a unit of measurement called “jícaras”, whose approximate volume is one litre. Each “jícara” of mushrooms has a price of 10 to 30 Mexican pesos, equivalent to USD$0.50 to USD$1.50 (Table 2). It is important to record that the prices of wild mushrooms have not changed significantly in the last 30 years (except for medicinal or entheogenic mushrooms). Additionally, cultivated Pleurotus sp. is sold at a price of up to 60 Mexican pesos per kilogram, equivalent to USD$3.00. However, people mentioned that wild mushrooms are of better quality and more nutritious compared to those cultivated. The commercialization of medicinal mushrooms is varied. The prices and the units to measure the number of mushrooms are diverse. They are sold by “jícaras”, pieces, sizes, in pairs or the enough amount to carry out a ritual ceremony, according to the species. The prices of medicinal mushrooms have varied significantly. For example, 30 years ago, the price for P. cubensis and P. caerulescens was 5 Mexican pesos, equivalent to USD$0.25. Nowadays, the price of these mushrooms varies according to their abundance, seasonality and vendor, but their average price is 600 Mexican pesos, equivalent to USD$30.00 (Table 2). Another major aspect of wild mushrooms trade is their commercialization by brokers or “coyotes”. These people are dedicated to commercialize manly Cantharellus spp. and Psilocybe spp.