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Traditional ecological knowledge (TEK) includes the understanding, innovations, and practices of indigenous and local communities throughout the world (Brush 1993, Berkes 1999). TEK is developed from experience over time, adapted to the local culture and environment, and is transmitted orally from generation to generation. It is collectively owned, and represented in stories, songs, folklore, cultural values, beliefs, rituals, community laws, local language and agricultural practices, including the production of plant and animal species (Berkes et al. 2000, CBD 2020). Traditional knowledge of bryophytes (mosses, liverworts, and hornworts) has been reported around the world but is often neglected (Berkes 1993, Harris 2008) in formal research. The study of the relationship between humans and these non-vascular plants is called ethnobryology (Flowers 1957). However, this term is not often used in ethnobiology, perhaps because there are few studies on bryophytes (Harris 2008). According to the last author, there are around 150 bryophyte species of ethnobotanical interest recorded worldwide. The highest number of reports have been made from such countries as China, India, the United States, and Canada; the most common ethnobotanical use for bryophytes is for medicinal purposes (Harris 2008, Alam 2016). Numerous examples are cited by Glime (2017b).
Mexico is one of the countries with the highest biological and cultural diversity in the world (Sarukhán et al. 2009, Toledo et al. 2019). About 25 % of the vascular plant species (approximately 7,461 species) are used in traditional ways (Mapes & Basurto 2016). Thus, the flora is an essential part of traditions and customs in many regions of the country. Nevertheless, bryophytes are usually neglected in ethnobotanical studies about traditional knowledge.
Recent estimations report 1,591 species of bryophytes living in different ecosystems in Mexico (Delgadillo 2014, Delgadillo & Juárez-Martínez 2014, Ellis et al. 2018, Escolástico-Ortíz & Juárez-Martínez 2018). Use of bryophytes has been documented since about 500 years for medicinal purposes and 180 years for religious purposes (De la Cruz 1552, Calderón de la Barca 1843); however, it is likely that they were also used in pre-Columbian times. Harris (2008) indicated that four species are used for handicrafts, and Delgadillo & Cárdenas (1990) reported the use of at least three moss species that are sold during the Christmas season. However, there is no consensus on which species are used and what their associated TEK is in different regions of Mexico. This study seeks to answer the following questions: how many bryophyte species are used by humans in Mexico? Is there any TEK associated with them? Specifically, the work intends 1) to obtain a checklist of useful bryophytes, including their use or associated TEK, and 2) to discuss the importance of these results for ethnobotany and for bryophyte conservation in Mexico.
Recording TEK associated with bryophytes updates knowledge of the use of non-timber forest resources, especially for a neglected group of plants. This work also seeks to stimulate the inclusion of bryophytes in ethnobotanical studies in Mexico, and in other countries with high biocultural diversity.
Materials and methods
The literature review searched for peer-reviewed articles and books in the Scopus and Google Scholar databases. Keywords included "bryophytes or moss or liverwort or hornwort" and "ethnobotany or traditional uses or traditional ecological knowledge or traditional knowledge or non-timber forest resources" (in English and Spanish) somewhere in the document. To obtain most records, all available works to date (2020) were considered. The Ethnobotanical Database of Plants of Mexico (BADEPLAM; Caballero & Cortés 1982-2020), the Digital Library of Traditional Mexican Medicine (Zolla & Argueta 2009), and the Herbarium of Medicinal Plants of the Mexican Institute of Social Security were reviewed. Also, personal records obtained between 2008 and 2017 in the state of Oaxaca, specifically in the municipalities of Ixtlán de Juárez (17° 19' 51" N, 96° 29' 15" W), Santiago Comaltepec (17° 33' 57" N, 96° 32' 54" W), and Miahuatlán de Porfirio Díaz (16° 19' 33" N, 96° 35' 31" W) were included. Personal records included species sampled for taxonomic determination; these were compared against specimens at MEXU.
The data were organized in a spreadsheet that included names of taxa, source document, and recorded locality. Local common names in native or indigenous languages were included, as well as an English and Spanish translation when possible. Traditional uses were arranged according to the classification by Bernal-Ramírez et al. (2019) which considers nine categories that are applicable to bryophytes: food, environmental, crafts, ceremonial, construction, fuel, medicinal, ornamental, and veterinary. Species sold during the Christmas season (Anastacio-Martínez et al. 2017a, b, Martínez-López et al. 2017), were classified in the "ceremonial" category.
Moss nomenclature was updated according to a revised version of LATMOSS (Delgadillo 2010) and for families based on Goffinet & Buck (2020); for liverworts and hornworts the classification accepted by Söderström et al. (2016) was followed. A TEK map showing the distribution of reports for bryophytes in Mexico was prepared.
Results
Few publications document TEK and use of bryophytes in Mexico (~30). Based on these references, the southern states contribute most records, particularly Oaxaca (15), Puebla (10), and Michoacán (8) (Figure 1). Mexican bryophytes with a use and TEK included 36 species in 30 genera and 23 families (Appendix 1, Figure 2). Literature on Mexican bryophytes does not report use as food, fuel, construction, or veterinary medicine. Mosses are the group with most uses (29), whereas liverworts only contributed 3 species. No references included information on hornworts.
Figure 1 Map of reports of useful bryophytes with associated TEK in different states of Mexico (black circles).
Figure 2 Some bryophytes species used in México with ceremonial (A. Campylopus sp., B. Dendropogonella rufescens, C. Dicranum sp., D. Meteorium sp., E. Pilotrichella flexilis, F. Thuidium delicatulum), craft (G. Bryum sp.), and medicinal (H. Marchantia sp., I. Pleurochaete squarrosa) proposes.
The most relevant use of bryophytes is ceremonial during the Christmas period, though it is not the only religious activity where they are used. The mosses Thuidium delicatulum (Hedw.) Schimp., Hypnum amabile (Mitt.) Hampe, Dendropogonella rufescens (Schimp.) E. Britton, and those in the genera Dicranum and Campylopus, were mostly used for this purpose over the last five decades. Other uses, including TEK, such as crafts, environmental, medicinal, and ornamental have also been documented (Appendix 1, Table 1). Results include native language names of species, but also generic names, to identify mosses as a group (Table 2). In some native languages, bryophytes are classified locally based on the substrates where they grow (Table 3). These languages include Huastec, Maya, Mixe, Nahuatl, Popoluca, Seri, Tarahumara, Totonac, Tzeltal, and Zapotec (Appendix 1, Tables 1-3).
Table 1 Traditional knowledge records that do not specify moss taxa. Information includes locality, native name (with the native language name), associated TEK description, and source. Use categories are represented by CE: Ceremonial, CR: Craft, EN: Environmental and ME: Medicinal.
| Locality | Native name | Use category and description | Reference |
|---|---|---|---|
| Center of Mexico | Tepitoton teamoxtli, Náhuatl | ME: Treatment of various illnesses by the Aztecs. | De la Cruz (1552) |
| Mexico City | CE: Used as decoration material to make the nativity scenes in post-independent Mexico. | Calderón de la Barca (1843) | |
| Center of Mexico | Amoxtli, Náhuatl. | CR: An aquatic moss (of unknown source) was used to make codices in pre-Hispanic times. This report is dubious and could include other organisms like algae. | Bay & Linares (2016) |
| San Pedrito (Otomi-Tepegua ethnicity), Hidalgo. | EN: Women of the community observed reduction in the availability of mosses in the forest. They record this knowledge because they are the group in charge of the use and management of the non-wood forest products. | Vázquez-García (2015) | |
| Las Brisas, Barra de Cahoacán, Chiapas. | EN: Mosses growing on bamboo (Otatea acuminata (Munro) C.E. Calderón & Soderstr.) are used as indicators of its maturity. This criterium is to choose those stems useful in the construction of rural housing. | Moreno-Martínez et al. (2012) | |
| North of Michocán state. | EN: The presence of mosses is considered essential for the establishment of agave plantlets (Agave inaequidens K. Koch) because it is related to adequate moisture conditions. This agave species is important in the local production of mezcal. | Torres et al. (2015) |
Table 2 Terms for “moss” in some native languages in Mexico.
| Native term | Language | Locality | Source |
|---|---|---|---|
| Con | Tzeltal | Chiapas state | Berlin et al. (1974) |
| Tecata | Spanish | Capécuaro, Michoacán | Delgadillo (2000) |
| Tsot-zé | Tzeltal | Lacanjá and laguna El Suspiro, Chiapas | Delgadillo (2000) |
| Tsa’nk | Mixe | Santa María Tlahuitoltepec, Oaxaca | Gallardo-Jiménez (2013) |
| Xibeu' | Zapotec | Miahuatlán District, Oaxaca | Ruegsegger & Ruegsegger (1955) |
| Mbaxhs* | Zapotec | Buenavista Loxicha, Oaxaca | Luna-José & Rendón-Aguilar (2012) |
| Yamasa* | Seri | Sonora state | Felger & Moser (1985) |
* According to the source, the term also applies to liverworts and lichens.
Table 3 Terms in native languages in Mexico to classify mosses according to the substrates where they grow.
| Native languages, localities, and source | ||||
|---|---|---|---|---|
| Substrate | Tzeltal, Chiapas (Berlin et al. 1974) | Náhuatl, Mexico (De la Cruz 1552) | Mayan, Lacanjá and laguna El Suspiro, Chiapas (Delgadillo 2000) | Zapotec, San Agustín Loxicha, Oaxaca (Luna-José & Rendón-Aguilar 2012) |
| Trees | Con teʔ = Tree moss |
|
||
| Rocks | Teamoxtli = Rock-inhabiting moss | Kushem-tunich = Attached to a rock | ||
| Soil | Con lumilal = Ground moss | Mbaxs lad ble = Growing on the ground | ||
Discussion
This study reports 36 species of bryophytes with their use or TEK. These results place Mexico among the countries with the highest number of ethnobryological records, third only to China (63 spp.) and India (58 spp.), and with a similar number as the United States (Harris 2008). Most reports come from areas with high biological diversity, humidity, and ethnic group's presence such as the states of Oaxaca and Chiapas. This relationship agrees with Toledo's (1990, 2019) suggestion regarding the presence of greater traditional knowledge in bio-culturally diverse areas. Species richness and lush growth in mild climates, however, are conditions that undoubtedly have favored and given ample opportunity for human interaction with potentially useful bryophyte species in certain areas of Mexico.
A traditional classification of mosses is reported for Chiapas, in Tzeltal and Maya languages, according to the substrate in which they grow (Berlin et al. 1974, Delgadillo 2000). In other regions of Mexico, there are traditional generic names that include liverworts and some lichens (Table 2) (Felger & Moser 1985, Luna-José & Rendón-Aguilar 2012). However, the translation of these terms refers to “moss” because it is the closest word to describe these cryptogams due to similarity in growth form, size, and appearance. Bryophytes may have different generic names within the same native language, possibly due to linguistic variations. The most evident cases of linguistic variations occur in Nahuatl, the language with the highest extant speaker number, and with the most variants in the country (Moseley 2007). Zapotec in Oaxaca state also has name variations, where the terms to describe mosses are different even between localities in the same region (Ruegsegger & Ruegsegger 1955, Luna-José & Rendón-Aguilar 2012).
Traditional moss uses related to Christmas or as decorative material are the most common. These results are in contrast with the worldwide trend of medicinal use that includes few records in Mexico (Alcorn 1984, Delgadillo 2000, Harris 2008, López-Santiago 2015, Glime 2017b). Using mosses to decorate Nativity scenes resulted from a Christmas tradition that began in the nineteenth century and has continued until today (Anastacio-Martínez et al. 2017b). Due to the large volumes harvested for this purpose, this represents a threat to bryophyte (moss) diversity and individual populations. On a national scale, the annual harvest can exceed 900 tons (Zamora-Martínez et al. 2008). Gómez-Peralta & Wolf (2001) documented the harvest of approximately 50 tons of bryophytes in the Monarch Butterfly Reserve in Michoacán state. Accordingly, 14 species were recorded for Christmas use in Mexico, and there may be more. However, this is not the only religious activity in which bryophytes are used. Other examples are the Patamban flower carpets in Michoacán state, as well as rituals in Zapotec communities in the northern Oaxaca state (Martínez-López et al. 2015, 2017, Cornejo-Tenorio & Ibarra-Manríquez 2019). For handicrafts, records of bryophytes included a dubious report on the manufacture of codex paper -making in ancient Mexico, and as filling material to lighten heavy loads by Tzeltales in Chiapas state (Berlin et al. 1974, Bay & Linares 2016).
Moss genera used for ceremonial and crafts purposes (e.g., Dicranum, Hypnum, Polytrichum, Thuidium) typically have intense colors, large size, abundance, and attractive growth forms (Anastacio-Martínez et al. 2017a, b, Cornejo-Tenorio & Ibarra-Manríquez 2019). These traits can be important for the development of productive and conservation activities of bryophytes (Grout 1934, Thieret 1956). In the Ixtlán de Juárez community, in Oaxaca state, these plants are part of the criteria to designate ecotourism areas, an important economic activity for the local population (Fuente-Carrasco & Ramos-Morales 2013). Bryophytes are the visual attraction of trails for visitors, where they form a large part of the plant biomass; this use promotes their conservation (pers. obs.). Thus, while some traditional practices represent a risk, others have positive effect on bryophyte conservation. Nevertheless, bryophytes used for medicinal and environmental purposes do not necessarily have aesthetic characteristics, suggesting an in-depth traditional knowledge of some of their phytochemical properties and ecological features. Some examples are the use of Sematophyllum adnatum to prepare medicinal teas in Yucatán (Glime 2017b), and small bryophyte species growing on the bamboo stem to estimate its maturity in Barra de Cahoacán, Chiapas (Moreno-Martínez et al. 2012).
De la Cruz's (1552) work can be considered as the oldest record of medicinal use of bryophytes in Mexico. However, there is a knowledge gap until new medicinal reports appeared in recent decades (Alcorn 1984, Delgadillo 2000, López-Santiago 2015, Glime 2017a). Bryophytes have a wide variety of secondary metabolites and they are used for antibiotics and pharmaceutical production (Pandey & Alam 2020). In Mexico, plants reported with antibacterial activity used in traditional medicine do not include bryophytes (Sharma et al. 2017). Therefore, the species presented in this review point to the necessity of such studies.
Regarding environmental uses, Torres et al. (2015) show that mosses are considered a suitable substrate for germination and seed development by farmers, of such species as Agave inaequidens K. Koch in Michoacán state. The capacity of bryophyte layers to promote seed germination has been previously reported; they reduce the risk of predation and maintain stable conditions of temperature and humidity necessary for their development (Tooren 1988, Soudzilovskaia et al. 2011). Another example of environmental use is cited by Jasso-Arriaga et al. (2016), where people conserve mosses layers in the forest floor to maintain the harvest of wild mushrooms in the state of Mexico. Other reports provide information on changes in bryophyte diversity in some regions of Mexico. For example, Vázquez-García (2015) shows that women in the community of San Pedrito, Hidalgo, observed a reduction in moss populations in the forest over time. Although it is not clear if the women knew the importance and roles of bryophytes in the forest, this information is a reference to possible effects of habitat degradation and climate change on a local scale (Wolverton et al. 2014). The environmental uses of bryophytes are among the rarest worldwide. Therefore, these represent an important antecedent of traditional ecological knowledge of bryophytes in Mexico thus far overlooked.
Mexico has an extensive heritage of useful plants, and bryophytes are part of this biocultural wealth. The information on the uses and traditional knowledge of bryophytes may increase with their inclusion in ethnobotanical works. It is not clear why bryophytes are not considered in these studies. Some reasons may include, for example, the assumption that bryophytes used in different parts of Mexico are the same species, the development of research limited to vascular flora, and the low value given by researchers to the identity of bryophyte species and their TEK (Kimmerer 2003). Collaboration between bryologists, ethnobiologists, and local people can improve this situation and avoid the risk of losing valuable information, especially when languages and traditional knowledge are at risk of disappearing (Moseley 2007, Campbell & Belew 2018).
This work provides a first list of the bryophyte’s species used in Mexico, as well as associated TEK. Thus, this study updates the information of useful bryophyte species in the country and highlights the ethnobotanical knowledge of a group of neglected plants.
