Some advances in the study of lichens since the time of E. M. Holmes

Some of the principal advances in our knowledge of lichens made since the time of E. M. Holmes are briefly reviewed. Floristic studies on British lichens have continued and distribution maps of particular species are now being produced. These studies have shown that many species have disappeared from areas affected by air pollution by sulphur dioxide. Lichen taxonomy has seen the acceptance of chemical characters as valid taxonomic criteria, and the realization that ascus types, ascocarp structure and ascocarp ontogeny are more important at the generic level than ascospore characters alone. Lichens are not now thought of as a single taxonomic group but as fungi related to various orders of non-lichenized Ascomycotina, united only in their common method of nutrition (symbiosis with algae); some lichenized and non-lichenized species with identical types of ascocarps are consequently being treated as members of the same genus. Considerable progress has been made in describing lichen communities characteristic of particular substrates and situations and in understanding the factors which may affect the occurrence of particular species. Lichens have proved to be particularly valuable indicators of mean air pollution levels and scales for the estimation of these levels based solely on the lichen vegetation have been devised. Physiological and ultrastructural studies on lichens have progressed rapidly in the last decade. Carbohydrates have been demonstrated to move from the algal to the fungal component and the mobile carbohydrates found to differ in lichens with different genera of algae. The algal and fungal partners have been isolated and studied in pure culture. Most attempts to synthesize lichens have met with little success but in 1970 a thallus was formed from its components in culture. It is now possible to grow some lichens under controlled conditions in the laboratory and these techniques, together with transplant experiments, will clearly assume considerable importance in the investigation of taxonomic and ecological problems in the future. Little progress has been made in the study of lichen cytology or the factors affecting the establishment of lichen thalli. Lichens constitute an important part of some ecosystems and their decline due to anthropogenic influences is consequently a matter of concern for all biologists.     

Diversity of Saxicolous Lichens along an Aridity Gradient in Central México

Lichens are symbiotic organisms that comprise a fungus and a photosynthetic partner wich are recognized as a good indicator of climate change. However, our understanding of how aridity affects the diversity of saxicolous lichens in drylands is still limited. To evaluate the relationship between saxicolous lichen diversity and aridity in a central México dryland, a geographical transect was established of 100 km to build an aridity gradient in the semiarid zone of the State of Querétaro, Mexico, comprising ten sampling sites with a 10 km separation. Species richness, abundance and diversity of soil lichen species were recorded using two sampling methods: the quadrat-intercept and the line-intercept method, to compare their performance in assessing soil lichen diversity in drylands. The number of species and Shannon diversity of saxicolous lichens were higher at intermediate values of the aridity index (AI = 0.10–0.34). Quadrat intercept and point intercept methods gave quite similar results, which means that the selected method does not influence the results in a significant way. This study confirms the role of saxicolous lichens as climate change indicators and reveals the importance of the sampling method selection in the evaluation of different parameters of soil lichen diversity in drylands.     

Lichen palatability depends on investments in herbivore defence

Lichens are well-suited organisms for experimental herbivory studies because their secondary compounds, assumed to deter grazing, can be non-destructively extracted. Thalli of 17 lichen species from various habitats were cut in two equal parts; compounds were extracted from one part by acetone, the other served as a control. These two pieces were offered as a paired choice to the generalist herbivore snail Cepaea hortensis. Control thalli of all lichens were consumed at a low rate regardless of their investments in acetone-extractable lichen compounds; naturally compound-deficient lichen species were not preferred compared to those with high contents. However, for extracted thalli, there was a highly significant positive correlation between rate of consumption and the extracted compound contents. These data imply that herbivore defence has evolved in different directions in different lichens. Studied members of Parmeliaceae, common in oligotrophic habitats, have high contents of carbon-rich acetone-soluble compounds; these lichens became highly palatable to snails subsequent to acetone rinsing. Extracted lichen compounds were applied to pieces of filter paper and fed to snails. Extracts from members of the Parmeliaceae significantly deterred feeding on paper. Such data suggest that generalist herbivores may have shaped evolution in the widespread and highly diverse Parmeliaceae towards high investments in lichen compounds. On the other hand, lichens belonging to the Physciaceae and Teloschistales, common in nutrient-enriched habitats, are deficient in, or have low concentrations of, lichen compounds. Such lichens did not become more palatable after acetone rinsing. The orange anthraquinone compound parietin, restricted to the Teloschistales, and which has previously been found to protect against excess light, did not deter grazing.     

新疆地衣研究现状与展望

地衣在自然界中是一种特殊的生物类群,具有独特的结构特征和较强的环境适应能力。它在环境污染评价,药物和香料开发利用方面具有较高的研究价值。目前对地衣的研究涉及地衣分类、地衣区系地理、地衣群落生态学、地衣化学、利用地衣评价环境质量以及地衣分子生物学等领域。回顾了新疆地衣研究近20年的发展概况,总结了新疆地衣研究的发展历程、特点,并展望了新疆地衣研究的方向。     

Degradation of the disease-associated prion protein by a serine protease from lichens

The disease-associated prion protein (PrP(TSE)), the probable etiological agent of the transmissible spongiform encephalopathies (TSEs), is resistant to degradation and can persist in the environment. Lichens, mutualistic symbioses containing fungi, algae, bacteria and occasionally cyanobacteria, are ubiquitous in the environment and have evolved unique biological activities allowing their survival in challenging ecological niches. We investigated PrP(TSE) inactivation by lichens and found acetone extracts of three lichen species (Parmelia sulcata, Cladonia rangiferina and Lobaria pulmonaria) have the ability to degrade prion protein (PrP) from TSE-infected hamsters, mice and deer. Immunoblots measuring PrP levels and protein misfolding cyclic amplification indicated at least two logs of reductions in PrP(TSE). Degradative activity was not found in closely related lichen species or in algae or a cyanobacterium that inhabit lichens. Degradation was blocked by Pefabloc SC, a serine protease inhibitor, but not inhibitors of other proteases or enzymes. Additionally, we found that PrP levels in PrP(TSE)-enriched preps or infected brain homogenates are also reduced following exposure to freshly-collected P. sulcata or an aqueous extract of the lichen. Our findings indicate that these lichen extracts efficiently degrade PrP(TSE) and suggest that some lichens could have potential to inactivate TSE infectivity on the landscape or be a source for agents to degrade prions. Further work to clone and characterize the protease, assess its effect on TSE infectivity and determine which organism or organisms present in lichens produce or influence the protease activity is warranted.     

Discoveries and identification methods of metal oxalates in lichens and their mineral associations: A review of past studies and analytical options for lichenologists

Lichens are generally acknowledged as the main agents of the initial bioweathering of rock substrates. The most direct evidence of a chemical interaction between lichens and their mineral substrata is the production of oxalic acid, of which interaction with ions present in a close lichen environment occasionally results in the precipitation of metal oxalates. In the past, only a few studies revealed the presence of metal oxalates in lichens, and currently, almost no new discoveries are arriving. Therefore, the main goal of this review is to bring the focus back to this phenomenon. To date, only Mg oxalate dihydrate, Mn oxalate dihydrate, Cu oxalate hemihydrate, Zn oxalate dihydrate, and anhydrous Pb oxalate have been detected in lichens. The most reliable diagnostic methods of metal oxalates in lichens are powder X-ray diffraction (pXRD), infrared spectroscopy (IR), extended X-ray absorption fine structure (EXAFS) spectroscopy, and Raman spectroscopy. Prospective lichen specimens for investigations may be found in anthropogenically-polluted environment or naturally metal-rich rock substrata. This review acts as an initial guide that provides analytical options for field lichenologists, offers a few suggestions for further research on this matter, and encourages to new biomineral discoveries in the scope of mineralogy. Survival in the metal-toxic environment, ability to produce oxygen, and extremophile nature, are the reasons why lichens are good subjects for research within the context of currently expanding astrobiology sciences, as well.     

Ecological and biotechnological aspects of lichens

Lichens and the partners from three different kingdoms are both taxonomically and physiologically a very diverse group, which makes them interesting from both ecological and biotechnological points of view. A lichen is a mutual ecophysiological innovation in many extreme environments in which symbiosis seems to protect the partners. Lichen’s ability to grow in harsh environments can be advantageous, resulting in important ecological niches, or disadvantageous when lichens occupy and cause biodeterioration of cultural monuments. Recently, new candidate compounds for drugs, UVB protection, and antifreeze proteins for frozen foods were discovered. Lichens were also found to have potential in bioplastic degradation and prevention of desertification. Nevertheless, there is still large potential for further industrial screening and research on lichen products. Due to improved culture techniques of isolated symbionts, increased knowledge of their secondary metabolism and improved methods for solubilizing lichen metabolites, the screening and activity tests can be implemented more easily today than in the past.     

Lichens as material for Lepidoptera's housing: A molecular approach to a widespread and highly selective interaction

Interactions between lichens and invertebrates are widespread and have evolved independently in several orders of invertebrates. Lichens participate as food, shelter, background for mimicry, or as camouflage for animals to wear. For this study, we discovered caterpillars of the moth family Psychidae living inside bags made from silk and lichens. We used molecular techniques to identify the lichens present and analyzed caterpillar selectivity for lichen species. We selected 13 bags and recovered the ITS genetic marker for every visually different lichen fragment. Obtained sequences were compared against a newly created ITS database from lichens of the study area. Caterpillars only used eight out of the 300 lichen species present showing a strong selection tendency for microfoliose lichens containing sekikaic acid. Our results suggest that caterpillars select lichen species at a higher rate than what is expected based on their local abundances. We provide an accessible way to study these widespread interactions in future projects.     

Lichen species richness is highest in non-intensively used grasslands promoting suitable microhabitats and low vascular plant competition

Lichens are very sensitive to habitat changes and their species richness is likely to decline under intensive land use. Currently, a comprehensive study analyzing lichen species richness in relation to land-use types, extending over different regions and including information on habitat variables, is missing for temperate grasslands. In three German regions we studied lichen species richness in 490 plots of 16 m² representing different land-use types, livestock types, and habitat variables. Due to the absence of low-intensity pastures and substrates such as woody plants, deadwood and stones, there were no lichens in the 78 plots in Schorfheide-Chorin. In the two other regions, the richness of lichen species was 45 % higher in pastures than in meadows, and 77 % higher than in mown pastures, respectively. Among the pastures, the richness of all lichen species was on average 10 times higher in sheep-grazed pastures than in the ones grazed by cattle or horses. On average, the richness of all lichen species increased by 3.3 species per additional microhabitat. Furthermore, the richness of corticolous lichens increased by 1.2 species with 10 % higher cover of woody plants, lignicolous lichen species richness increased by 4.8 species with 1 % higher cover of deadwood, and saxicolous lichen species richness increased by 1.0 species with 1 % higher cover of stones. Our findings highlight the importance of low-intensity land use for lichen conservation. In particular, the degradation of grasslands rich in microhabitats and the destruction of lichen substrates by intensification, and conversion of unfertilized pastures formerly grazed at low intensity to meadows should be avoided to maintain lichen diversity.     

Diversity and distribution of terricolous lichens as indicator of habitat heterogeneity and grazing induced trampling in a temperate-alpine shrub and meadow

Lichens are among the most sensitive biomonitors of ecosystem health and human induced disturbances. Terricolous lichens of Chopta–Tungnath (Garhwal, western Himalaya, India) were analysed for their ability to indicate habitat variability and disturbances induced by livestock grazing. Terricolous lichens were sampled from 12 sites, distributed across the three macrohabitats between 2,700 and 4,001 m, using 50 × 10 cm narrow frequency grids having five 10 × 10 cm sampling units. The terricolous lichen community of the area constituted, 20 species belonging to 10 genera, five families and four growth forms. Altitude and relative humidity were the major habitat factors found influencing the terricolous lichen community of the landscape. Fruticose and compound soil lichen growth forms were found indicative of habitat disturbance largely caused by grazing induced trampling. Terricolous lichen diversity of the area was delimited by grazing pressure at mid-altitudes (3,000–3,400 m) and by decreasing soil cover at higher altitudes (>3,400 m).     

Nutrient scavenging activity and antagonistic factors of non-photobiont lichen-associated bacteria: a review

Lichens are defined as the specific symbiotic structure comprising a fungus and a green alga and/or cyanobacterium. Up until recently, non-photobiont endothallic bacteria, while known to be present in large numbers, have generally been dismissed as functionally irrelevant cohabitants of the lichen thallus, or even environmental contaminants. Recent analyses of lichen metagenomes and innovative co-culture experiments have uncovered a functionally complex community that appears to contribute to a healthy lichen thallus in several ways. Lichen-associated bacteriomes are typically dominated by several lineages of Proteobacteria, some of which may be specific for lichen species. Recent work has implicated members of these lineages in several important ecophysiological roles. These include nutrient scavenging, including mobilization of iron and phosphate, nitrogen fixation, cellulase, xylanase and amylase activities, and oxidation of recalcitrant compounds, e.g. aromatics and aliphatics. Production of volatile organic compounds, conferring antibacterial and antifungal activity, has also been demonstrated for several lichen-associated isolates. In the present paper we review the nature of non-phototrophic endolichenic bacteria associated with lichens, and give insight into the current state of knowledge on their importance the lichen symbiotic association.     

Extremotolerance and Resistance of Lichens: Comparative Studies on Five Species Used in Astrobiological Research II. Secondary Lichen Compounds

Lichens, which are symbioses of a fungus and one or two photoautotrophs, frequently tolerate extreme environmental conditions. This makes them valuable model systems in astrobiological research to fathom the limits and limitations of eukaryotic symbioses. Various studies demonstrated the high resistance of selected extremotolerant lichens towards extreme, non-terrestrial abiotic factors including space exposure, hypervelocity impact simulations as well as space and Martian parameter simulations. This study focusses on the diverse set of secondary lichen compounds (SLCs) that act as photo- and UVR-protective substances. Five lichen species used in present-day astrobiological research were compared: Buellia frigida, Circinaria gyrosa, Rhizocarpon geographicum, Xanthoria elegans, and Pleopsidium chlorophanum. Detailed investigation of secondary substances including photosynthetic pigments was performed for whole lichen thalli but also for axenically cultivated mycobionts and photobionts by methods of UV/VIS-spectrophotometry and two types of high performance liquid chromatography (HPLC). Additionally, a set of chemical tests is presented to confirm the formation of melanic compounds in lichen and mycobiont samples. All investigated lichens reveal various sets of SLCs, except C. gyrosa where only melanin was putatively identified. Such studies will help to assess the contribution of SLCs on lichen extremotolerance, to understand the adaptation of lichens to prevalent abiotic stressors of the respective habitat, and to form a basis for interpreting recent and future astrobiological experiments. As most of the identified SLCs demonstrated a high capacity in absorbing UVR, they may also explain the high resistance of lichens towards non-terrestrial UVR.     

石质文物地衣生物腐蚀及防治的研究进展

本文主要从生物因素方面考虑,分析了导致石质文物病害的主要原因和机理,并综述了国内外石质文物保护的现状、防治和修复方法。地衣是引起石质文物腐蚀的关键生物类群,与国外相比,目前国内专门针对地衣进行石质文物防治的研究鲜见报道。作者建议,石质文物的地衣生物腐蚀及防治研究应从以下方面入手：（1）石质文物地衣必须首先进行系统的种类调查;（2）对地衣腐蚀石质文物的机理进行深入研究;（3）探寻地衣病害安全有效的去除方法;（4）逐步探索地衣病害修复后的监测和预防方法。     

Epiphytic lichen synusiae and functional trait groups in boreo‐nemoral deciduous forests are influenced by host tree and environmental factors

Deciduous forests with temperate broad‐leaved tree species are particularily important in terms of biodiversity and its protection, but are threatened habitats in northern Europe. Using multivariate analyses we studied the effect of forest site type, environmental variables and host tree properties on epiphytic lichen synusiae as well as on the composition of species‐specific functional traits. Epiphytic lichens were examined on Acer platanoides, Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra and U. laevis in two types of forests: Humulus‐type floodplain forests and Lunaria‐type boreo‐nemoral forests on the talus slopes of limestone escarpment (klint forests). Klint forests located near the seashore were under greater maritime influence compared to floodplain forests located in inland Estonia which experience stronger air temperature contrasts. In addition to stand level and climatic variables, tree level factors (bark pH, trunk circumference and cover of bryophytes) considerably affected the species composition of the lichen synusiae. Overall, 137 lichen species were recorded, including 14 red‐listed species characteristic of deciduous trees. We defined 13 lichen societies and showed their preference to forests of a specific site type and/or host tree properties. In forests of both types, most of the epiphytic lichens were crustose, and had apothecia as the fruit bodies and chlorococcoid algae as the photobiont. However, the proportion of lichens with a foliose or fruticose growth form, as well as the proportion of lichens with vegatative diaspores, were higher in floodplain forests. In klint forests with a stronger influence from the wind, crustose species completely dominated, while species with vegetative diaspores were rare and most species dispersed sexually. Lichens with Trentepohlia as the photobiont were characteristic of these forests, and lichens with lirellate ascomata were prevailing, indicating the great uniqueness of the kint forests for epiphytic lichens in the boreo‐nemoral region.     

新疆石人沟山区岩面生地衣群落分布格局的环境解释

地衣作为真菌和绿藻/蓝绿藻的成功共生体,广泛分布在陆地生态中的各种栖息地。岩面生地衣作为陆地生态系统的主要组成部分,在干旱和半干旱地区陆地食物链中具有重要地位,同时对岩石的生物腐蚀和土壤的形成有重要作用。岩面生地衣的多样性和分布格局强烈地受到海拔、湿度、温度、降水量、太阳辐射强度和基物的特征(岩石类型、岩石大小、岩石的化学成分和营养成分)等多种因素的影响。为了研究乌鲁木齐县石人沟山区岩面生地衣群落与基物间的关系,该研究在乌鲁木齐县石人沟山区设立16个样地,计测样地中岩面生地衣的盖度,包括坡度、坡向、光照强度等7个环境因子,采用典范对应分析法(CCA)对各群落的物种分布格局与环境因子的关系进行了探讨。结果表明:石人沟山区的岩面生地衣共有27种,隶属于7目9科15属。其中,黄枝衣目、茶渍目和鸡皮衣目的种类较多,占该地区岩面生地衣总数的74.07%。CCA排序结果显示坡度、坡向、光照强度、湿度、岩石pH值是5个影响岩面生地衣种类分布格局的主要环境因子,并显示了岩面生地衣与样地间的对应性。     

Flechten. Der Erfolg einer Symbiose

The success of a symbiosis: Lichens Lichens are a unique group of organisms composed of one or two alga and a fungus. Together they form species specific thalli. Their common eco‐physiological properties allow colonizing almost all terrestrial habitats, even the most hostile climatic zones on earth. However, as poikilohydrous organisms they also suffer from disadvantages related with their nature. As water content cannot be actively controlled, many lichens experience water‐oversaturation, thus being not able to gain full photosynthetic rates, even though they have otherwise optimal conditions. These eco‐physiological properties set up the frame for which microclimatic situation the realized thallus construction might do best. As all optimizations regarding water uptake also count for water loss, lichens are always at the edge of having either too much or not enough water for optimal carbon gain. So each habitat has its own challenge for the lichen thallus construction and lichens have to fit well into a specific ecological niche.     

How lichens impact on terrestrial community and ecosystem properties

Lichens occur in most terrestrial ecosystems; they are often present as minor contributors, but in some forests, drylands and tundras they can make up most of the ground layer biomass. As such, lichens dominate approximately 8% of the Earth's land surface. Despite their potential importance in driving ecosystem biogeochemistry, the influence of lichens on community processes and ecosystem functioning have attracted relatively little attention. Here, we review the role of lichens in terrestrial ecosystems and draw attention to the important, but often overlooked role of lichens as determinants of ecological processes. We start by assessing characteristics that vary among lichens and that may be important in determining their ecological role; these include their growth form, the types of photobionts that they contain, their key functional traits, their water‐holding capacity, their colour, and the levels of secondary compounds in their thalli. We then assess how these differences among lichens influence their impacts on ecosystem and community processes. As such, we consider the consequences of these differences for determining the impacts of lichens on ecosystem nutrient inputs and fluxes, on the loss of mass and nutrients during lichen thallus decomposition, and on the role of lichenivorous invertebrates in moderating decomposition. We then consider how differences among lichens impact on their interactions with consumer organisms that utilize lichen thalli, and that range in size from microfauna (for which the primary role of lichens is habitat provision) to large mammals (for which lichens are primarily a food source). We then address how differences among lichens impact on plants, through for example increasing nutrient inputs and availability during primary succession, and serving as a filter for plant seedling establishment. Finally we identify areas in need of further work for better understanding the role of lichens in terrestrial ecosystems. These include understanding how the high intraspecific trait variation that characterizes many lichens impacts on community assembly processes and ecosystem functioning, how multiple species mixtures of lichens affect the key community‐ and ecosystem‐level processes that they drive, the extent to which lichens in early succession influence vascular plant succession and ecosystem development in the longer term, and how global change drivers may impact on ecosystem functioning through altering the functional composition of lichen communities.     

Does secondary chemistry enable lichens to grow on iron-rich substrates?

Lichen substances are shown to increase or to inhibit the adsorption of Fe at cation exchange sites. The influence on the adsorption strongly differs between individual lichen substances and is different for Fe²⁺ and Fe³⁺. These results add a new biological role to the known functions of lichen secondary metabolites. In an experiment with cellulose filters, which were soaked with acetone solutions of lichen substances and were then incubated with micromolar solutions of FeCl₂ or FeCl₃, many lichen substances were found to increase Fe³⁺ adsorption, whereas others had no effect. Most lichen substances had no effect on Fe²⁺ adsorption, but two were found to reduce and one to increase the level of adsorption. Lichens of Fe-poor and -rich sites contain lichen substances with different adsorption behavior towards Fe²⁺ and Fe³⁺. All the studied lichen substances, which only occur in lichens of Fe-poor sites, turned out to be effective Fe³⁺ adsorbents. Lichens of Fe-bearing rock and slag, however, were found to lack lichen substances, or to contain substances that did not adsorb Fe³⁺ and had no effect on Fe²⁺ adsorption, or thirdly, to contain substances that increased Fe³⁺ adsorption, but decreased Fe²⁺ adsorption. These results suggest that lichen substances do play a significant role in Fe adsorption in lichens and determine their tolerance to excess concentrations of Fe. Notwithstanding the strong correlation between the secondary chemistry of lichen species and their preference for Fe-rich or Fe-poor substrates, the postulated mechanism of temporary Fe adsorption by lichen substances has to be subject of future biochemical research.     

Ice encapsulation protects rather than disturbs the freezing lichen

Arctic and alpine terricolous lichens are adapted to harsh environments and are tolerant to extremely low temperatures when metabolically inactive. However, there are reports indicating that freezing can be lethal to metabolically active lichens. With a projected warmer and more unstable climate, winter precipitation at high latitudes will fall more frequently as rain, causing snowmelt and encapsulating terricolous lichens in ice or exposing them to large temperature fluctuations. Lichens are a major winter food source for reindeer in most parts of the circumpolar region. A laboratory experiment tested how three hydrated reindeer forage lichen species covered by snow, encapsulated in ice, or uncovered responded to storage at freezing temperatures and subsequent warming. Photosynthetic performance (maximal fluorescence of dark-adapted samples and net photosynthetic rates) was significantly lower in lichens not insulated by snow or ice, whereas there were few differences between the snow and ice treatments. It is suggested that snow and ice provide sufficiently moist environments to improve extracellular and reduce intracellular ice nucleation activity. Ice encapsulation, which is often lethal to vascular plants, did not have any negative effects on the studied lichens. The results indicate that complete snow and ice melt followed by refreezing can be detrimental to terricolous lichen ecosystems. Reduced lichen biomass will have a negative effect both on reindeer winter survival and the indigenous peoples who herd reindeer.     

中国地衣的濒危等级评估

地衣是大型真菌的重要组成部分, 是中国大型真菌红色名录评估的主要对象之一。本次评估的地衣共2,164种, 其中子囊菌地衣2,145种, 担子菌地衣19种, 分属于2门9纲34目92科352属。评估结果显示, 地衣受威胁物种共28种, 占被评估地衣物种总数的1.29%, 其中极危3种, 濒危7种, 易危18种; 近危6种, 无危657种, 而数据不足的物种达1,473种。这类因为缺乏足够的数据资料而未能进一步评估其生存状态和红色名录等级的物种占评估地衣总数的68.07%, 凸显了我国地衣学研究的严峻形势及对地衣分类学人才的迫切需要。对受威胁物种的分析表明, 地衣由于自身生长缓慢和对环境污染敏感, 并且绝大多数分布区域狭窄, 种群数量少, 对生境退化的适应和恢复生长的能力弱, 人类活动和气候变化导致的栖息地破坏和碎片化使得这些地衣的生存受到严重威胁。值得注意的是, 评为易危的一些地衣, 因具有较大的食药用价值而被过度采挖利用, 如不采取有效保护措施, 其受威胁程度将进一步加剧。