diversity Review Ethnolichenology—The Use of Lichens in the Himalayas and Southwestern Parts of China Mei-Xia Yang 1,2, * , Shiva Devkota 3,4 , Li-Song Wang 5 and Christoph Scheidegger 1,2, * 1 2 3 4 5 * Swiss Federal Institute for Forest, Snow and Landscape Research WSL, 8903 Birmensdorf, Switzerland Faculty of Sciences, University of Bern, 3012 Bern, Switzerland Global Institute for Interdisciplinary Studies (GIIS), Kathmandu 3084, Nepal; shiva.devkota@gmail.com Himalayan Climate & Science Institute (HCSI), Washington, DC 20007, USA Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, CAS, Kunming 650201, China; wanglisong@mail.kib.ac.cn Correspondence: meixia124@gmail.com (M.-X.Y.); christoph.scheidegger@wsl.ch (C.S.); Tel.: +41-79-836-8441 (M.-X.Y.); +41-79-460-7132 (C.S.) Abstract: Lichens are used in traditional medicine, food and various other ethnic uses by cultures across the Himalayas and southwestern parts of China. Evidence-based knowledge from historical and modern literatures and investigation of ethnic uses from 1990 proved that lichen species used as medicine in the Himalayas and southwestern parts of China totaled to 142 species; furthermore, 42 species were utilized as food. Moreover, some lichens are popularly used for lichen produce in ethnic and modern life. An understanding and clarification of the use of lichens in the Himalayas and southeastern parts of China can therefore be important for understanding uses of lichens elsewhere and a reference for additional research of lichen uses in the future.



 Keywords: lichen; ethnic use; medicinal; edible species; Himalayas; southwestern China Citation: Yang, M.-X.; Devkota, S.; Wang, L.-S.; Scheidegger, C. Ethnolichenology—The Use of Lichens in the Himalayas and Southwestern 1. General Introduction of Lichen Uses Parts of China. Diversity 2021, 13, 330. Lichens are composite organisms containing algae (e.g., Trebouxia or Trentepohlia), or cyanobacteria (Nostoc), living among filaments of multiple fungal species in a mutualistic relationship [1,2]. Lichens dominate vegetation types on about 7% of the planet’s surface; additionally, they are important components of primary producers in a wide range of substrates and habitats, including some of the most extreme conditions on earth (North and South Pole, desert, even glass surfaces, etc.) [3]. Many lichens (such as Usnea Dill. ex Adans., Everni Ach., Hypogymnia (Nyl.) Nyl., Parmelia Ach. et al.) are very sensitive to environmental disturbances and they can be used to assess air pollution [4–6]. Unlike simple dehydration in plants and animals, lichens may experience a complete loss of body water in dry periods [7]. Lichens are important in contributing nitrogen to soils either by forming litter, or predation by herbivores, e.g., snails, which then defecate, providing nitrogen to the soils [8]. In deserts and semi-arid areas, lichens are part of extensive, living biological soil crusts, essential for maintaining the soil structure. They have a long fossil record in soils, dating back 2.2 billion years [9]. Lichens are also important in diets for humans and animals. Based on research on the diet of Rhinopithecus roxellana Milne-Edwards in China, lichens are the most eaten food for Rh. roxellana (Figure 1a), accounting for 38.4% of the overall diet [10]. The regional diversity of lichens will also affect the changes in the living area of Rh. roxellana. Moreover, for the human diet, mostly in the temperate and arctic regions of the world, people usually use lichens as food, pharmaceutical products, and various ethnic uses [11]. In the past, Iceland moss (Cetraria islandica (L.) Ach.) was an important source of food for humans in northern Europe and the lichen was cooked as bread, porridge, pudding, soup, or salad. Wila (Bryoria fremontii (Tuck.) Brodo & D. Hawksw.) was an important food in parts of North America, where it was usually pit cooked. Northern peoples in North https://doi.org/10.3390/d13070330 Academic Editors: Edit Farkas and Michael Wink Received: 9 June 2021 Accepted: 14 July 2021 Published: 18 July 2021 Publisher’s Note: MDPI stays neutral with regard to jurisdictional claims in published maps and institutional affiliations. Copyright: © 2021 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (https:// creativecommons.org/licenses/by/ 4.0/). Diversity 2021, 13, 330. https://doi.org/10.3390/d13070330 https://www.mdpi.com/journal/diversity Diversity 2021, 13, 330 Diversity 2021, 13, 330 2 of 15 2 of 16 Wila (Bryoria fremontii (Tuck.) Brodo & D. Hawksw.) was an important food in parts of North America, where it was usually Northern peoples in North America and Siberia traditionally eatpit thecooked. partially digested reindeer lichenAmerica (Cladoniaand P. Siberia traditionally eat the partially digested reindeer lichen (Cladonia P. Browne) after Browne) after they remove it from the rumen of caribou or reindeer that have been killed. they tripe remove it from the rumen of Lasallia caribouMérat) or reindeer that have been killed. Rock tripe Rock (Umbilicaria Hoffm. and is a lichen (sometimes more species of (Umbilicaria Hoffm. and Lasallia Mérat) is a lichen (sometimes more species of lichens) that lichens) that has frequently been used as an emergency food in North America and one has frequently beenesculenta used as (Miyoshi) an emergency food North and one species, Umspecies, Umbilicaria Minks, is in used in aAmerica variety of traditional Korean bilicaria esculenta (Miyoshi) Minks, is used in a variety of traditional Korean and Japanese and Japanese foods [12]. foods [12]. Figure Lichensare areused usedbyby animals and humans. (a) Rhinopithecus roxellana eating in Figure1. 1. Lichens animals and humans. (a) Rhinopithecus roxellana eating UsneaUsnea in southsouthwestern China; (b) Bai minority people harvesting lichens in Yunnan; (c) Ethnic lichen marwestern China; (b) Bai minority people harvesting lichens in Yunnan; (c) Ethnic lichen market in Yunnan. (d) Freshly gathered lichens for the kitchen Nepal; in (e)Nepal; Health-promoting tea products ket in Yunnan. (d) Freshly gathered lichens for theinkitchen (e) Health-promoting teaof Lethariella and Thamnolia; (a–c,e) photographed by Li-Song Wang; (d) photographed by Shiva Devproducts of Lethariella and Thamnolia; (a–c,e) photographed by Li-Song Wang; (d) photographed by kota. Devkota. Shiva Ethnolichenology is is aa branch branch of of Ethnolichenology of ethnobotany ethnobotanythat thatstudies studiesthe theuses usesthat thatman manmakes makes lichens traditionally [13,14]. Lichens areare used forfor many different medicinal purposes, but Lichens used many different medicinal purposes, of lichens traditionally [13,14]. there are are some general categories of use that that reoccur across the world. Lichens are often but there some general categories of use reoccur across the world. Lichens are often as a decoction to ailments treat ailments relating to either the or lungs the digestive drunkdrunk as a decoction to treat relating to either the lungs the or digestive system system is particularly common the Himalayas and southeastern [15–18].[15–18]. This is This particularly common in the in Himalayas and southeastern China.China. Many Many otherofuses of lichens are related to treating gynecological may be other uses lichens are related to treating gynecological diseases.diseases. This mayThis be related to related to the common use of lichens for treating sexually transmitted infections and the common use of lichens for treating sexually transmitted infections and aliments of the aliments of the [11]. urinary [11].of Two other of lichens that less common, urinary system Twosystem other uses lichens thatuses are less common, butare reoccur in several but reoccur in several cultures, areafflictions the treatment of eye afflictions and use[11]. in different cultures, are different the treatment of eye and use in smoking mixtures smoking mixtures Besides, lichens are usedwounds, externally for as dressing wounds, Besides, lichens are[11]. often used externally foroften dressing either a disinfectant or either asbleeding a disinfectant or to stop bleeding [18]. Other common lichen uses for to stop [18]. Other common topical lichen uses are fortopical skin infections andare sores, skin infections and sores, including sores in the mouth [11,19]. Many of the traditional Diversity 2021, 13, 330 3 of 16 medicinal uses of lichens are probably related to their secondary metabolites, many of which are known to both be physiologically active and act as antibiotics [15]. However, some of the traditional uses of lichens also rely on the qualities of lichen carbohydrates. Many of the traditional uses of lichens involve boiling the lichen to create a mucilage which is drunk for lung or digestive ailments, or applied topically for other issues [17,18,20]. Other lichen carbohydrates which may be important are the isolichenins and galactomannans, which are widespread across various taxonomic groups of lichens, and the pustulins, that are found in Umbilicariaceae [11]. Today, ethnic groups inhabiting the mighty Himalayas (Bhutan, China, India and Nepal) primarily adapt the classical systems of medicine following Ayurveda, Siddha, Unani, Traditional Chinese Medicine (TCM) and Amchi practices and continue their traditional uses of lichens for food, beverages and traditional medicine. Within the last decade, however, the sale of lichens for folk uses, especially for supposedly health-promoting teas, has increased remarkably, as the Himalayas and southwestern parts of China have become popular regions for domestic tourism (Figure 1b,e). Since 1990, the authors have studied these folk uses as part of a broader investigation of the lichen flora of southwestern parts of China, with aims to reveal the species diversity of lichens used traditionally and currently, the diversity of various uses by studying records from herbaria and other notes and interviewing current population. The present paper summarizes the results of our ethnobotanical investigation, which has already been treated in our previous studies [21–26], and other related research references, which are listed in this study. We expect that the understanding of the use of lichens in the Himalayas and southeastern parts of China will serve as an important reference for additional research of lichen uses. 2. Materials and Methods Chinese samples are available in the Lichen Herbarium of the Kunming Institute of Botany (KUN-L) and we reviewed useful lichen collections housed at different herbaria (KATH, TUCH) and at the Natural History Museum, Tribhuvun University, Nepal, for their additional notes, if any. We interviewed local people about their uses of lichens and visited ethnic markets to buy samples of the lichens offered for sale and also to ensure the legitimacy, relevance and credibility of the given evidence. Specimens were examined using standard microscopic techniques and hand-sectioned under a Nikon SMZ 745T dissecting microscope. Anatomical descriptions are based on observations of these preparations under a Nikon Eclipse 50i microscope. Secondary metabolites of all the specimens were identified using spot tests and thin-layer chromatography (TLC), as described by White and James [27] and Orange et al. [28]. Solvent system C (toluene:acetic acid = 85:15) was used for TLC analysis. 3. Results 3.1. Traditional Medicinal Lichens in the Himalayas and Southwestern Parts of China Introduction of Typical Medicinal Lichens Some of the typical, traditionally applied lichens are presented in Figure 2. Several species of Lobaria (Schreb.) Hoffm. are frequently used traditional medicines effective for treating pneumonia in the Himalayas and southwestern China, due to their lung-like appearance (applied because of the doctrine of signatures, suggesting that herbs can treat body parts that they physically resemble) [23,29]. Lobaria species have also been reported to serve as a valuable source of proteins, having a protein content higher than that of kelp or edible fungi, such as Tremella Pers. In addition, the content of dietary fiber in Lobaria species is significantly higher than in other fungi and edible algae and Lobaria species are rich in calcium [30]. Similarly, Peltigera leucophlebia (Nyl.) Gyeln. is used as a supposed cure for thrush (Aphtha, Candidiasis), due to the resemblance of its cephalodia to the appearance of the disease [31]. The earliest report about traditional medicin of lichen could be the Usnea longissima Ach. in the Chinese Qing dynasty (ca. 1500); it was reported that the Diversity 2021, 13, 330 Diversity 2021, 13, 330 4 of 15 4 of 16 appearance of the disease [31]. The earliest report about traditional medicin of lichen could be the Usnea longissima Ach. in the Chinese Qing dynasty (ca. 1500); it was reported that the Usnea species were used as litmus, to treat cold, swelling and pain [32]. Lichens Usnea species were used litmus, to treat cold, swelling pain [32]. Lichens also have also have their place in as current pharmaceutical research;and lichens produce metabolites of their place in current pharmaceutical research; lichens produce metabolites of potential potential therapeutic or diagnostic value [18]. Some metabolites produced by lichens are therapeutic diagnostic valuesimilar [18]. Some metabolites produced by lichens structurally structurallyorand functionally to broad-spectrum antibiotics, whileare a few of them and functionally similar to broad-spectrum antibiotics, while a few of them are are associated, respectively, with antiseptic similarities [33]. Usnic acid is the associated, most comrespectively, with antiseptic similarities [33]. Usnic is the mostresearch commonly monly studied metabolite produced by lichens [15]. acid It is also under as astudied bacterimetabolite produced by lichens [15]. It is also under research as a bactericidal agent against cidal agent against Escherichia coli (Migula) Castellani & Chalmers and Staphylococcus auEscherichia coli (Migula) Castellani & Chalmers and Staphylococcus aureus Bergey. reus Bergey. Figure2.2. Selected Selected traditional traditional medicinal medicinal lichens. Figure lichens. (a) (a) Sulcaria Sulcariasulcata sulcata(Lév.) (Lév.)Bystrek; Bystrek;(b) (b)Cladonia Cladoniasp.; sp.; (c) Lobaria sp.; (d) Usnea longissima. Photographed by Li-Song Wang. (c) Lobaria sp.; (d) Usnea longissima. Photographed by Li-Song Wang. 3.2. 3.2. Summary Summary of of Research Researchon onMedicinal MedicinalLichens Lichens Textual Textualresearch researchof ofseveral severalpublications publicationshave havedocumented documentedthe theinventory inventoryand andethnic ethnic uses uses of of lichens lichens from from Nepal Nepal [22,34–37], [22,34–37], India India [20–41], [20–41],Bhutan Bhutan [19,42] [19,42]and andsouthwestern southwestern parts parts of of China, China, in inancient ancient[32] [32]and andmodern modernresearch research[14,23,25,43,44]. [14,23,25,43,44].The Theliterature literatureand and investigation of folk usages proved that lichen species used as medicine in the Himalayas investigation of folk usages proved that lichen species used as medicine in the Himalayas and China totaled totaled to to142 142species speciesbelonging belongingtoto1616families families and and southwestern southwestern parts of China and 46 46 genera. Figure providesthe thespecies speciesnumber numberwithin within the the different genera genera. Figure 3 3provides genera of of traditional traditional medicinal medicinal lichens. lichens. Table Table 11presents presentslichen lichenspecies speciesin inalphabetical alphabeticalorder orderand andprovides providesthe the details detailson oneach eachtraditional traditionaluse. use. Diversity2021, 2021,13, 13,330 330 Diversity 16 5 5ofof15 Figure3.3.Species Speciesnumber numberwithin withinthe thedifferent differentgenera generaof oftraditional traditionalmedicinal medicinallichen. lichen. Figure Lichenspecies speciesused usedin intraditional traditionalmedicine medicineor orherbal herbalmedicine medicineininthe theHimalayas Himalayasand andsouthwestern southwesternparts partsofofChina. China. Table 1. 1.Lichen Table The newly added ones through our investigation are indicated in boldface. The newly added ones through our investigation are indicated in boldface. Current Species Names Current Species Names Alectoria ochroleuca (Schrank) A. Massal. Anzia formosana Asahina Alectoria ochroleuca (Schrank) A. Massal. Anzia japonica (Tuck.) Müll. Anzia formosana AsahinaArg. AnziaAnzia japonica (Tuck.) Müll. Arg. opuntiella Müll. Arg. Anzia Müll.Asahina Arg. Anzia opuntiella ornata (Zahlbr.) Anzia ornata (Zahlbr.) Asahina Bryoria asiatica (Du Rietz) Brodo & D. Hawksw. Bryoria asiatica (Du Rietz) Brodo & D. Hawksw. Bryoria bicolor (Hoffm.) Brodo & D. Hawksw. Bryoria bicolor (Hoffm.) Brodo & D. Hawksw. Bryoria confusa (D.D. Awasthi) Brodo & D. Hawksw. Bryoria confusa (D.D. Awasthi) Brodo & D. Cetraria sp. Hawksw. Cetraria sp. (L.) Ach. Cetraria islandica Cetraria islandica (L.) Ach. Cetraria laevigata Rass. Cetraria laevigata Rass. Cetrariella delisei (Bory ex Schaer.) Kärnefelt & Cetrariella delisei (Bory Schaer.) Kärnefelt & A. ex Thell A. Thell Cetrelia cetrarioides (Delise) W.L. Culb. & C.F. Culb. Cetrelia cetrarioides (Delise) W.L. Culb. & C.F. Cetrelia olivetorum (Nyl.) W.L. Culb. & C.F. Culb. Culb. Cetrelia pseudolivetorum (Asahina) W.L. Culb. & Cetrelia olivetorum (Nyl.) W.L. Culb. & C.F. Culb. C.F. Culb. Cetrelia pseudolivetorum (Asahina) W.L. Culb. & C.F. Culb. Function and Function Traditional and TradiApplication tional Apa, b plication a, b b a b a a a a a l l l l l l a a a a a a Folk Names Folk Names 树发 树发 (Tree hair) （Tree hair） 树发 树发 (Tree hair) （Tree hair） 树发 (Tree hair) 树发 （Tree hair） Main Area of Main Area Use of Use References References China [16,43] China China China China China China China China China [43] [16,43] [43] [43] [43] [29] [29] [29,43] [29,43] China China China China China China India India China China China China China China [23] [23] [23] [23] [23] [23] [17,45] [17,45] [23] [23] [23] [23] [43] [43] a, b China [43] a, b a, b China China [43] [43] a, b China China [29] [43] China [29] a a Diversity 2021, 13, 330 6 of 16 Table 1. Cont. Current Species Names Function and Traditional Application Folk Names Main Area of Use References Cladia aggregata (Sw.) Nyl. b 石花菜 (Stone flower) China [29,43] Cladonia amaurocraea (Flörke) Schaer. a, b 青雪茶 (Green snow-tea) China [16] Cladonia arbuscula (Wallr.) Flot. l China [16] Cladonia bellidiflora (Ach.) Schaer. a China [16] Cladonia cenotea (Ach.) Schaer. b China [29,43] Cladonia cervicornis (Ach.) Flot. c China [16,46] Cladonia crispata (Ach.) Flot. o India [17,20] Cladonia cyanipes (Sommerf.) Nyl. a China [29,43] Cladonia digitata (L.) Hoffm. d China [16] Cladonia fenestralis Nuno l China [23] Cladonia fruticulosa Kremp. b, d China [16,43] Cladonia floerkeana (Fr.) Flörke a China [43] Cladonia gracilis (L.) Willd. c China [23,47,48] Cladonia macilenta Hoffm. a, b China [43] Cladonia macroceras (Delise) Ahti l China [29] Cladonia mitis Sandst. a China [16,43] Cladonia pleurota (Flörke) Schaer. a China [29,43] 太白鹿角 (Taibai antlers) Cladonia pyxidata (L.) Hoffm. b China [43] Cladonia rangiferina (L.) Weber b China [29] Cladonia squamosa (Scop.) Hoffm. b China [29,43] Cladonia stellaris (Opiz) Pouzar & Vězda a 太白花 (Tai-bai flower) China [23,29,43] Dolichousnea diffracta (Vain.) Articus (as Usnea diffracta Vain.) k 老君须 (Lao Jun’s beard) China [23,47] Dolichousnea longissima (Ach.) Articus (as Usnea longissima Ach.) a, n, q 松萝/老君须 (Lao Jun’s beard) China, India [18,23,38] Dolichousnea trichodeoides (Vain. ex Motyka) Articus (as Usnea trichodeoides Vain.) a, b China [29] Dendriscosticta wrightii (Tuck.) B. Moncada & Lücking (as Sticta wrightii Tuck.) d China [46,49] Dermatocarpon miniatum (L.) W. Mann d, e China [50] Eumitria baileyi Stirt. (as Usnea baileyi (Stirt.) Zahlbr.) m India [18] Eumitria pectinata (Taylor) Articus (as Usnea pectinata Taylor) a, b China [29] Evernia divaricata (L.) Ach. a, f China [29,43] Evernia esorediosa (Müll. Arg.) Du Rietz a, f, g China [29,43] Evernia mesomorpha Nyl. a, f China [29,43] Flavoparmelia caperata (L.) Hale a China [16,29] Flavopunctelia flaventior (Stirt.) Hale a, b China [16,43] Flavopunctelia soredica (Nyl.) Hale a, b China [43] Diversity 2021, 13, 330 7 of 16 Table 1. Cont. Current Species Names Function and Traditional Application Heterodermia comosa (Eschw.) Follmann & Redón b Heterodermia diademata (Taylor) D.D. Awasthi b, d Heterodermia hypochraea (Vain.) Swinscow & Krog Main Area of Use References China [16,43] China, India, Nepal [16,17,22,43] b China [29,43] Heterodermia pseudospeciosa (Kurok.) W.L. Culb. b China [29,43] Heterodermia speciosa (Wulfen) Trevis. b China [16] Hypogymnia flavida McCune & Obermayer a China [29,43] Hypogymnia hypotrypa (Nyl.) Rass. a China [29,43] Hypogymnia physodes (L.) Nyl. b China [29,43] Hypogymnia pseudoenteromorpha M.J. Lai b China [43] Hypotrachyna cirrhata (Fr.) Divakar, A. Crespo, Sipman, Elix & Lumbsch (as Everniastrum cirrhatum (Fr.) Hale) f China [23,51,52] Hypotrachyna nepalensis (Taylor) Divakar, A. Crespo, Sipman, Elix & Lumbsch a, f China, Nepal [16,23,37] Hypotrachyna sinuosa (Sm.) Hale a China [29,43] Leptogium delavayi Hue l China [23] Leptogium saturninum (Dicks.) Nyl. l China [23] Leptogium trichophorum Müll. Arg. l China [23] Lethariella cladonioides (Nyl.) Krog b 红雪茶 红雪茶 (Red snow-tea) China [24,29] Lethariella flexuosa (Nyl.) J.C. Wei h 红雪茶 红雪茶 (Red snow-tea) China [24] Lethariella zahlbruckneri (Du Rietz) Krog h 红雪茶 红雪茶 (Red snow-tea) China [24,44] Leucodermia boryi (Fée) Kalb (as Heterodermia boryi (Fée) Kr.P. Singh & S.R. Singh) b China [16,43] Lobaria sp. o Bhutan [19] Lobaria isidiosa (Müll. Arg.) Vain. e 老龙皮 老龙皮 (Dragon skin) China [23,29] Lobaria kurokawae Yoshim. e 树蝴蝶 树蝴蝶 (Tree butterfly) China [29] Lobaria meridionalis Vain. e 树蝴蝶 树蝴蝶 (Tree butterfly) China [29] Lobaria orientalis (Asahina) Yoshim. l 树蝴蝶 树蝴蝶 (Tree butterfly) China, India [17,23,45] Lobaria pindarensis Räsänen l 树蝴蝶 树蝴蝶 (Tree butterfly) China [23] Lobaria pulmonaria (L.) Hoffm. d 蛤蟆七 蛤蟆七 (Toad skin) China [17,48] Lobaria retigera (Bory) Trevis. d, i 老龙皮 (Dragon skin) China [23,30] Lobaria sublaevis (Nyl.) Yoshim. d 树蝴蝶 (Tree butterfly) China [43] Folk Names झुलो (Jhulo) याउ (Jhyauu) 老龙皮 树蝴蝶 树蝴蝶 Diversity 2021, 13, 330 8 of 16 Table 1. Cont. Current Species Names Function and Traditional Application Folk Names Main Area of Use References Lobaria yunnanensis Yoshim. l 树蝴蝶 (Tree butterfly) China [23] Mycoblastus alpinus (Fr.) Th. Fr. ex Hellb. c China [16,46,50] Myelochroa irrugans (Nyl.) Elix & Hale b China [24] Nephromopsis cucullata (Bellardi) Divakar, A. Crespo & Lumbsch (as Flavocetraria cucullata (Bellardi) Kärnefelt & A. Thell) a China [43] Nephromopsis nivalis (L.) Divakar, A. Crespo & Lumbsch (as Flavocetraria nivalis (L.) Kärnefelt & A. Thell) a China [29] Nephromopsis pallescens (Schaer.) Y.S. Park c China [23] Omphalodina chrysoleuca (Sm.) S.Y. Kondr., L. Lőkös & Farkas (as Rhizoplaca chrysoleuca (Sm.) Zopf) d China [16] Ophioparma lapponica (Räsänen) Hafellner & R.W. Rogers c China [23,29] Ophioparma ventosa (L.) Norman c China [23,29] Oxneria fallax (Arnold) S.Y. Kondr. & Kärnefelt (as Xanthoria fallax Arnold) b China [16,29] Parmelia adaugescens Nyl. b China [16] Parmelia saxatilis (L.) Ach. a, b China [23,29,43] Parmelia sulcata Taylor b China [29,43] Parmelina quercina (Willd.) Hale b China [29,43] Parmelinella wallichiana (Taylor) Elix & Hale (as Parmelina wallichiana (Taylor) Hale) b China [43] Parmotrema abessinicum (Nyl. ex Kremp.) Hale a India [17] Parmotrema cetratum (Ach.) Hale (as Rimelia cetrata (Ach.) Hale & A. Fletcher) b China [16] Parmotrema chinense (Osbeck) Hale & Ahti a India [45] Parmotrema nilgherrense (Nyl.) Hale a, d, e, l India [45] Parmotrema reticulatum (Taylor) M. Choisy (as Rimelia reticulata (Taylor) Hale & A. Fletcher) b China [29,43] Parmotrema sancti-angelii (Lynge) Hale p India [17,20,38] Parmotrema subtinctorium (Zahlbr.) Hale c China [16] Parmotrema tinctorum (Despr. ex Nyl.) Hale a, b China [29,43] Peltigera aphthosa (L.) Willd. d China [16,29] Peltigera canina (L.) Willd. j China [16,45] Peltigera polydactylon (Neck.) Hoffm. l China, India [16,17,29] Punctelia borreri (Turner) Krog a, b China [16] Ramalia sp. d Jhyauu (Unnecessary stuff) Nepal [22] Ramalina commixta Asahina f 石花菜 (Stone flower) China [51,52] 石花 (Rock flower) Rathipuvvu (Rock flower) Chharila Jhau Diversity 2021, 13, 330 9 of 16 Table 1. Cont. Current Species Names Function and Traditional Application Folk Names Main Area of Use References Ramalina conduplicans Vain. d 石花菜 (Stone flower) China [16] Ramalina fastigiata (Pers.) Ach. a, f 石花菜 (Stone flower) China [29,43] Ramalina roesleri (Schaer.) Nyl. b, f 石花菜 (Stone flower) China [29,43] Ramalina sinensis Jatta a, f 石花菜 (Stone flower) China [29,43] Stereocaulon exutum Nyl. c China [29] India [17] Dhungo-ku-Jhau (Rock flower) Stereocaulon himalayense D.D. Awasthi & I.M. Lamb Stereocaulon japonicum Th. Fr. b, d China [23] Stereocaulon myriocarpum Th. Fr. b China [23] Stereocaulon paschale (L.) Hoffm. b China [23,29] Stereocaulon tomentosum Fr. b, d China [23,43] India [17] China [43,51,52] 石寄生 (Rock parasite) Sticta gracilis (Müll. Arg.) Zahlbr. 石花菜 (Stone flower) Sulcaria sulcata (Lév.) Bystrek f, h Sulcaria virens (Gyeln.) Bystrek c China [23] Teloschistes flavicans (Sw.) Norman l China [16] Thamnolia subuliformis (Ehrh.) W.L. Culb. k 白雪茶 (White snow-tea) China [23,24,43] Thamnolia vermicularis (Sw.) Schaer. k 白雪茶 (White snow-tea) China [23,24,32,43] Umbilicaria esculenta (Miyoshi) Minks d, h 石耳 (Stone ear) China [23,29,43] Umbilicaria hypococcinea (Jatta) Llano a China [16,29] Umbilicaria nanella Frey & Poelt d China [48,53] Umbilicaria vellea (L.) Ach. l China [16] Umbilicaria yunnana (Nyl.) Hue l China [23] Usnea aciculifera Vain. h China [16] Usnea ceratina Ach. k China [16] Usnea florida (L.) F.H. Wigg. a, c China [43,47,50] Usnea nidifica Taylor l China [16] Usnea rubicunda Stirt. a China [29,43] India [17,20] Usnea rubrotincta Stirt. Usnea subfloridana Stirt. a China [23] Usnea subsordida Stirt. r India [19,54] Usnea spp. a, d, i, n India, Bhutan [17,18,20,42] Varicellaria velata (Turner) I. Schmitt & Lumbsch (as Pertusaria velata (Turner) Nyl.) c China [23,29] Diversity 2021, 13, 330 10 of 16 Table 1. Cont. Current Species Names Function and Traditional Application Vulpicida juniperinus (L.) J.-E. Mattsson & M.J. Lai Main Area of Use References a China [43] Vulpicida pinastri (Scop.) J.-E. Mattsson & M.J. Lai a China [43] Xanthoparmelia camtschadalis (Ach.) Hale a China [16,29] Xanthoparmelia taractica (Kremp.) Hale a China [29] Xanthoparmelia tinctina (Maheu & A. Gillet) Hale a China [29] Xanthoria parietina (L.) Th. Fr. b, d China [16,29,43] Folk Names The abbreviations indicate different functions and traditional applications: “a”, raw material for antibiotics; “b”, raw material for making litmus reagent; “c”, hemostatic of external injury; “d”, anti-inflammatory and antibacterial; “e”, digestion facilitator and stomach enhancer; “f”, raw material for making spice; “g”, antihypertensive; “h”, anti-colon cancer; “i”, calcium level booster; “j”, treatment of rabies and icterus; “k”, detoxifying and cough suppressant; “l”, other medicinal; “m”, mixed with other aromatic herbs, such as Valeriana jatamansi, for flavor and curing tobacco; “n”, against lung troubles, hemorrhages and asthma attacks, strengthening of hair, treatment of skin eruptions and boils, stopping nose bleeds, preventing or treating blisters; “o”, anti-tumor; “p”, against skin diseases; “q”, treatment of bone fracture; “r”, flavoring tobacco. 3.3. Edible Lichens in the Himalayas and Southwestern Parts of China 3.3.1. Introduction of Typically Edible Lichen In the past, edible lichens were mostly gathered for private consumption [24]. In recent years, edible lichens are increasingly sold by local people or tourists as a commodity, after drying in the Himalayas and southwestern parts of China (Figure 4). Lethariella (Motyka) Krog and Thamnolia Ach. ex Schaer. are widely used as health-promoting teas, Lobaria, Umbilicaria Hoffm., Nephromopsis Müll. Arg. and Ramalina Ach. are used as food and are relatively common in the local supermarkets and some restaurants. Normally, summer and autumn are the best seasons to harvest edible lichens, which are used fresh or dried for later use. Usually, stewing with burns, steaming, cooking soup and other methods are used to make dishes, such as “Liang Ban” Ramalina, etc. 3.3.2. Summary of Research on Edible Lichen By researching ancient and modern literature and investigating folk usages in the Himalayas and southwestern parts of China, it is obvious that people generally name edible lichen “shuhua” or “shihuacai”; the names mean flowering form of an organism growing on the trees or stones and these phrases are still in use. Here we document a total of 42 species belonging to 18 genera of lichens that are edible (Table 2). The most commonly used genera of lichens are foliose or fruticose growth forms (such as Hypotrachyna, Lethariella, Lobaria, Nephromopsis, Ramalina, Thamnolia and Umbilicaria). The edible lichens counted in this article are limited and more investigation and research are needed in the future. Diversity 2021, 13, 330 Lethariella (Motyka) Krog and Thamnolia Ach. ex Schaer. are widely used as promoting teas, Lobaria, Umbilicaria Hoffm., Nephromopsis Müll. Arg. and Ramal are used as food and are relatively common in the local supermarkets an restaurants. Normally, summer and autumn are the best seasons to harvest edible 11 of 16 which are used fresh or dried for later use. Usually, stewing with burns, steaming, soup and other methods are used to make dishes, such as “Liang Ban” Ramalina, Figure 4. Selected ediblein lichens in the and southwestern China. (a) Lo Figure 4. Selected edible lichens the Himalayas and Himalayas southwestern China. (a) Lobaria; (b) Scrambled Scrambled eggs with Lobaria; (d) “Liang Ban” Ramalina; eggs with Lobaria; (c) Ramalina fastigiata;(c) (d) Ramalina “Liang Ban”fastigiata; Ramalina; (e) Nephromopsis pallescens (Schaer.) (e) Nep Y.S. Park; (f) Egg custard with (g) Lethariella; (h) Lethariella tea. pallescens; Photographed pallescens (Schaer.) Y.S. Nephromopsis Park; (f) pallescens; Egg custard with Nephromopsis (g) Letha by Li-Song Wang. Lethariella tea. Photographed by Li-Song Wang. Diversity 2021, 13, 330 12 of 16 Table 2. Edible lichens in Himalayas and southwestern China. Consumed Species Name Bryoria asiatica (Du Rietz) Brodo & D. Hawksw. Bryoria confusa (D.D. Awasthi) Brodo & D. Hawksw. Cetraria laevigata Rass. Cladia aggregata (Sw.) Nyl. Cladonia gracilis (L.) Willd. Cladonia stellaris (Opiz) Pouzar & Vězda Dermatocarpon miniatum (L.) W. Mann Hypotrachyna cirrhata (Fr.) Divakar, A. Crespo, Sipman, Elix & Lumbsch Hypotrachyna nepalensis (Taylor) Divakar, A. Crespo, Sipman, Elix & Lumbsch Leptogium delavayi Hue Leptogium saturninum (Dicks.) Nyl. Leptogium trichophorum Müll. Arg. Leptogium wilsonii Zahlbr. (as Leptogium denticulatum Nyl.) Leucodermia leucomelos (L.) Kalb (as Heterodermia leucomelos (L.) Poelt) Lobaria isidiophora Yoshim. Lobaria isidiosa (Müll. Arg.) Vain. Lobaria kurokawae Yoshim. (Eaten) as vegetable, with different cooking methods, such as stewing, steaming, boiling, frying, making soup, etc. [17,18,21–24,26,30,37] Lobaria orientalis (Asahina) Yoshim. Lobaria pindarensis Räsänen Lobaria pulmonaria (L.) Hoffm. Lobaria retigera (Bory) Trevis. Lobaria yunnanensis Yoshim. Nephromopsis pallescens (Schaer.) Y.S. Park Parmotrema cetratum (Ach.) Hale Parmotrema reticulatum (Taylor) M. Choisy Parmotrema tinctorum (Despr. ex Nyl.) Hale Ramalina commixta Asahina Ramalina conduplicans Vain. Ramalina fastigiata (Pers.) Ach. Ramalina roesleri (Schaer.) Nyl. Ramalina sinensis Jatta Sulcaria sulcata (Lév.) Bystrek Umbilicaria esculenta (Miyoshi) Minks Umbilicaria hypococcinea (Jatta) Llano Umbilicaria yunnana (Nyl.) Hue Usnea longissima Ach. Diversity 2021, 13, 330 Diversity 2021, 13, 330 12 of 15 13 of 16 Consumed (Drunk) as tea [24,26,30] Umbilicaria esculenta (Miyoshi) Minks Table 2. Cont. Umbilicaria hypococcinea (Jatta) Llano Umbilicaria yunnana (Nyl.) Hue Species Name Usnea longissima Ach. Cladonia fenestralis Nuno Cladonia fenestralis Nuno Lethariella cladonioides (Nyl.)cladonioides Krog Lethariella (Nyl.) Krog Lethariella flexuosa (Nyl.) J.C. Wei (Nyl.) J.C. Wei Lethariella flexuosa (Drunk) as tea [24,26,30] Lethariella zahlbruckneri (Duzahlbruckneri Rietz) Krog (Du Rietz) Krog Lethariella Thamnolia subuliformis Thamnolia subuliformis (Ehrh.) W.L. Culb.(Ehrh.) W.L. Culb. Thamnolia (Sw.) Schaer. Thamnolia vermicularis (Sw.) vermicularis Schaer. Other Ethnic Ethnicand andModern ModernUses Uses 4.4. Other Lichens are are mainly mainly used used by byhumans humans for formedicine medicineand andfoods foodsin inthe theHimalayas Himalayasand and Lichens southwestern parts parts of of China, China, but but we we have have also also found found many many other, other,novel noveluses usesfor forthese these southwestern organisms in a local place. Lethariella, only distributed at around 3700–4300 m in the organisms in a local place. Lethariella, only distributed at around 3700–4300 m in the Himalayas, is mainly sold in Yunnan and Shangri-La, China, and also exported to Taiwan Himalayas, is mainly sold in Yunnan and Shangri-La, China, and also exported to Taiwan andJapan. Japan.InInthethe Himalayas, Tibet is a holy of Buddhism and a great and Himalayas, Tibet is a holy place place of Buddhism and there is athere greatisdemand demand for Tibetan incense; Lethariella is also used as an important component of Tibetan for Tibetan incense; Lethariella is also used as an important component of Tibetan incense incense of because of its special(Figure fragrance Besides, in currentofapplications of because its special fragrance 5a). (Figure Besides,5a). in current applications lichens, Usnea lichens, Usnea Sulcaria arematerials used for raw materialsand of perfume in Yunnan and Sulcaria areand used for raw of perfume fragranceand in fragrance Yunnan (Figure 5b). (Figure 5b). Cladonia isused commonly useddecoration as garden in decoration in China.etDevkota et al. [22] Cladonia is commonly as garden China. Devkota al. [22] reported reported ritual and spiritual value (RSV), aesthetic and decorative value (ADV), bedding ritual and spiritual value (RSV), aesthetic and decorative value (ADV), bedding value (BV) value (BV) and ethno-veterinary values (EVV) of lichens, together with medicinal and ethno-veterinary values (EVV) of lichens, together with medicinal value (MV)value and (MV)value and (FV), food among value (FV), among different andpeople indigenous people and local food different collectors andcollectors indigenous and local communities communities (IPLCs) in Nepal. CetreliaW.L. collata (Nyl.) W.L. Culb. & C.F.name: Culb.Platysma (current (IPLCs) in Nepal. Cetrelia collata (Nyl.) Culb. & C.F. Culb. (current name: Platysma is used fiber, as a sacrificial Melanelia infumata collatum Nyl.) iscollatum used asNyl.) a sacrificial together fiber, with together Melaneliawith infumata (Nyl.) Essl., (Nyl.) Essl.,cirrhatum Everniastrum cirrhatum nilgherrense, and Parmotrema nilgherrense, Usnea ghattensis G. Everniastrum and Parmotrema Usnea ghattensis G. Awasthi, for coloring Awasthi, for coloring hair, Thamnolia vermicularis (Sw.) Schaer., with its spiritual value in hair, Thamnolia vermicularis (Sw.) Schaer., with its spiritual value in India and Nepal [22,45], India and subsororioides, Nepal [22,45],used and to Buellia color palms and lips as a and Buellia colorsubsororioides, palms and lipsused as a to substitute for Heena, mostly substitute for Heena, mostly the Garhwali Uttarakhand and India [39]. by the Garhwali Herdsman in by Uttarakhand andHerdsman India [39]. in Furthermore, Shukla et al. [55] have also highlighted theal.use of have eleven lichen species as dying agents in lichen Gharwal region Furthermore, Shukla et [55] also highlighted the use of eleven species as of India. dying agents in Gharwal region of India. Figure5.5. (a) (a) Lethariella, Lethariella, used used for for Tibetan Tibetan incense; incense; (b) Figure (b) Usnea Usnea and and Sulcaria, Sulcaria, used usedfor forraw rawmaterials materialsofof perfume in Yunnan; (a) Photographed by Mei-Xia Yang; (b) photographed by Li-Song Wang. perfume in Yunnan; (a) Photographed by Mei-Xia Yang; (b) photographed by Li-Song Wang. Discussion and andConclusions Conclusions 5.5. Discussion Ourinvestigation investigationand andsurvey surveyof ofthe theliterature literatureindicate indicatethat that142 142lichen lichenspecies speciesare areused used Our as asmedicine medicineand and42 42species speciesare areused usedas asfood foodin inthe theHimalayas Himalayasand andsouthwestern southwesternparts partsof of China. considerable overlap between the medicinal and consumed lichens;lichens; except China.We Wefound found considerable overlap between the medicinal and consumed for three species of edible lichens (Leptogium wilsonii, Leucodermia leucomelos and Lobaria Diversity 2021, 13, 330 14 of 16 isidiophora), other species with edible uses also have medicinal functions (Tables 1 and 2). Therefore, the popularity of consuming healthy food items might be explained by their preventive role. Almost all food lichens are cooked in some way before being eaten and the cooking process is often complex, usually involving steps to remove toxins from the lichen [11,22,23,25,43]. For the medicinal lichens, the secondary compounds and carbohydrates are useful to humans. The studies reported that the nutritionally relevant carbohydrates in lichens include the glucans lichenin and isolichenin [11,14]. Some lichens also have significant levels of proteins and essential amino acids, as well as some minerals and vitamins, but most lichens only have minimal amounts of these nutrients [11,12,15,30]. Some lichens are only eaten in times of famine, some are a staple food or even a delicacy in the Himalayas and southwestern China. The medicinal and edible usage of lichen as healthy food is becoming more and more popular among local people and tourists. However, two obstacles are often encountered when eating lichens: lichen polysaccharides are generally indigestible to humans and lichens usually contain mildly toxic secondary compounds that should be removed before eating. Very few lichens are poisonous, but those having high concentrations of vulpinic acid or usnic acid are toxic [56]. Lichen resources are especially abundant in the Himalayas and southeastern parts of China and it is also a major advantage that they mostly grow at high altitudes without human activity and pollution. In recent years, resource survey and research on lichens as food and medicine have also been reported in these areas. Unfortunately, there are still some problems concerning the classification of species and the unclear distribution of resources; therefore, for example, the species classification of the genus Lobaria and the resource distribution of Lobaria in these areas also need to be investigated. At present, most lichens for food and traditional medicine are used directly as lichen raw materials and, considering that many lichens grow slowly, it results that lichens productivity is usually low. However, there are still some ecosystems of foliose lichens (e.g., Lobaria pulmonaria) that can produce significant lichen biomass within approximately a decade [57,58]. The low productivity of lichens means that over-harvesting is a real concern. For example, the genus Lethariella—with known distribution only in the Himalayas and southeastern parts of China—has reached an endangered state before its active ingredients and mechanisms could be fully understood. Based on our recent study, we expect to carry out further relevant research on effective medicinal and nutritional ingredients of lichens in the future and explore ways to obtain the required effective ingredients through artificial culturing or by fermentation, also to reduce the dependence on natural resources. Therefore, combining results of lichen taxonomy, ecology, chemistry and pharmacology is a top priority for our forthcoming research. At the same time, effective protection measures for some endangered species are important for the sustainable use of lichen resources. Author Contributions: Conceptualization, C.S. and M.-X.Y.; methodology, M.-X.Y.; software, M.-X.Y.; validation, M.-X.Y. and C.S.; formal analysis, M.-X.Y.; investigation, L.-S.W., C.S. and S.D.; resources, L.-S.W. and S.D.; data curation, M.-X.Y., L.-S.W. and S.D.; writing—original draft preparation, M.X.Y.; writing—review and editing, M.-X.Y., C.S., S.D. and L.-S.W.; visualization, M.-X.Y. and C.S.; supervision, C.S.; project administration, C.S.; funding acquisition, C.S., S.D. and L.-S.W. All authors have read and agreed to the published version of the manuscript. Funding: This work was supported by the Swiss National Science Foundation (grant JRP IZ70Z0_ 131338/1 to C.S.), the National Natural Science Foundation of China (No. 31970022, 31670028), the Second Tibetan Plateau Scientific Expedition and Research Program (No. 2019QZKK0503), the Global Biodiversity Information Facility/Biodiversity Fund for Asia (Project No: BIFA5_023 to S.D.) and the China Scholarship Council (CSC No. 201704910901). Institutional Review Board Statement: Not applicable. Data Availability Statement: The data that support the findings of this study are available from the corresponding author, upon reasonable request. Diversity 2021, 13, 330 15 of 16 Acknowledgments: We sincerely thank Xin-Yu Wang (Kunming Institute of Botany, CAS, China) and Dong Liu (Central South University of Forestry and Technology, China), for supporting expedition and research in China; the Lichen Herbarium of the Kunming Institute of Botany, CAS for providing research support; Ram Prasad Chaudhary (RECAST Kathmandu, Nepal) and Krishna Kumar Shrestha (Tribhuvan Univ. Kathmandu, Nepal), for supporting research in Nepal; and Karma Tshering, for giving insights into ethnobotany in Bhutan. Conflicts of Interest: The authors declare no conflict of interest. References 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 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