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Mycobionts of many lichen genera appear to demonstrate strong selectivity in the choice of algal partner. The biological properties of a photobiont and its availability in an environment significantly determine the habitat requirements of lichens. Flexibility in photobiont choice extends the ecological amplitude of lichens; therefore, it may constitute an important adaptive strategy for colonization of extreme habitats. The photobiont inventory of the three epigeic lichens most resistant to soil pollution, i.e., Cladonia cariosa, C. rei, and the hyperaccumulator Diploschistes muscorum, was examined to verify whether and to what extent algal composition depends on the type of habitat and substrate enrichment with heavy metals. Photobionts Asterochloris and Trebouxia were identified in the studied lichen species; however, the presence of Trebouxia was directly related to anthropogenic sites with technogenic substrates, and the proportion of lichen specimens with these algae clearly depended on the level of heavy-metal soil pollution and the habitat type. The total number of algal haplotypes increased with increasing soil pollution, and the richness was associated more with soil pollution than with a given lichen species. Additionally, a large number of lichen individuals bearing multiple algal genotypes at polluted sites were recorded. Although Cladonia lichens were previously thought to be restricted to Asterochloris, they are able to start the relichenization process with Trebouxia under specific habitat conditions and to establish a stable association with these algae when colonization of disturbed sites takes place. Comparative analysis of the internal transcribed spacer (ITS) rDNA sequences revealed as many as 13 haplotypes of Trebouxia, and phylogenetic analysis grouped them into two different clades. Such a high level of genetic diversity indicates that Trebouxia is well adapted to metal pollution and could be an alternative photosynthetic partner for certain lichens, especially in polluted sites.

Heavy metals present in the environment can cause a variety of injury symptoms in various organisms including lichens. Most studies examined metal-induced stress under controlled laboratory conditions, and little is known about actual response of lichens in their natural habitat. This study aims to recognize the effect of heavy metal accumulation (total and intracellular) on lichen physiological and biochemical parameters specifically related to the functioning of fungal component. Cladonia rei was used as a model species due to its common occurrence both in unpolluted and extremely polluted sites. We observed a decline in the fungal metabolism which was expressed by a decrease in ergosterol content and an increase in cell membrane damage as a result of increased Zn, Cd, Cu and Ni accumulation. Additionally, the results indicated that increased accumulation of xenobiotics (Pb and As) caused reduction of glutathione (GSH) concentrations and increased membrane lipid peroxidation. Therefore, we conclude that GSH does not provide high oxidative stress protection in C. rei which is somewhat against its insensitivity to pollution. The reduced pool of GSH could be explained by its oxidation to glutathione disulphide induced by heavy metal stress or its use for phytochelatin (PC) synthesis. The content of secondary metabolites was not related to heavy metal accumulation and remained at a relatively stable level. This indicates that the decline in the physi-ological condition did not weaken the mycobiont of C. rei enough to inhibit the synthesis of secondary metabolites and their precursors were supplied at a sufficient level. Thus, the potential function of main secondary metabolites as extracellular metal immobilizers and antioxidants is still possible even in individuals growing at extremely polluted sites. Despite the evident heavy metal stress, C. rei copes well and spreads easily through extremely polluted environments, which underlines its unique pioneering abilities in highly disturbed sites.

The synthesis of various unique secondary metabolites by lichens is the result of mutualistic symbiotic association between the mycobiont and autotrophic photobiont. The function of these compounds and causal factors for their production are not fully understood. This paper examines the effect of heavy-metal bioaccumulation and physiological parameters related to photosynthesis and carbon metabolism on the production of lichen substances in hyperaccumulator Diploschistes muscorum. The obtained model of secondary metabolite concentrations in the thalli demonstrates that the carbon source provided by the photobiont and associated polyols produced by the mycobiont have positive impact on the production; on the contrary, the increased intracellular load of heavy metals and excessive loss of cell membrane integrity adversely affected secondary metabolite contents. Additionally, the production of secondary metabolites appears to be more dependent on intracellular metal concentrations than on soil pollution level. To compensate for metal stress, both efficient functioning of algal component and sufficient production of secondary metabolites are required. The balanced physiological functioning of mycobiont and photobiont constitutes the complex protective mechanism to alleviate the harmful effects of heavy metal stress on primary and secondary metabolism of lichens.

The data presented here are related to the research article entitled “Cryptogamic communities as a useful bioindication tool for estimating the degree of soil pollution with heavy metals” (Rola and Osyczka, 2018) [1]. These data concern the relationships between epigeic cryptogamic biota and heavy metal concentrations in soil of areas associated with Zn–Pb industry. The presence of particular species and coverage of lichens and bryophytes as well as soil chemical parameters in relation to three different soil pollution classes and five habitat types are provided. Included data could be used to compare cryptogamic community structure and pollutant concentration levels with other Zn–Pb polluted areas. Keywords: Lichens, Bryophytes, Cryptogamic biota structure, Zn–Pb ores, Post-industrial areas, Environmental assessment

Phenotypic traits of lichens can be greatly modified by environmental factors. Granulose thalli on soil and podetia, densely covered with granules, referring to common and widespread lichen Cladonia cervicornis subsp. verticillata were found near zinc smelter. The granules are stratified, filled with fungal medulla and heavily encrusted with calcium oxalate weddellite crystals, not observed on regularly developed thalli of the species. Phylogenetic analysis revealed that deformed granulose forms belong to this taxon, showing that the phenotypic plasticity of the lichens of Cladonia can lead to the emergence of features that do not coincide with the taxonomic definition of the species. The heavy-metal accumulation capacity of both granulose and regular form of primary and secondary lichen thallus, in relation to the element content in corresponding substrate, was determined. Granulose-modified thalli accumulate greater amounts of heavy metals than regular ones, meaning that the bioaccumulation property of a given species may be greatly affected by morphological modifications. The granulose forms are also characterised by considerably higher ratios of Cd, Pb and As concentrations in lichen samples in relation to the corresponding substrates than regular ones. This means that collection of variously formed thalli should be avoided in biomonitoring sampling procedures. The results indicate that a substantial part of the element load, in particular zinc, in the examined lichen thalli collected near the smelter originates from atmospheric fallout. Electronic supplementary material The online version of this article (10.1007/s11356-017-0639-1) contains supplementary material, which is available to authorized users.

Heavy metals are known for their negative impact on the physiological processes of lichen photobiont. In spite of this, certain lichens are known to be effective pioneers of polluted sites. Cladonia cariosa, C. rei, and Diploschistes muscorum are prominent examples of lichens that spontaneously colonise post-industrial wastes. We examined the effect of total and intracellular Zn, Pb, Cd, As, Cu, and Ni accumulation in the thalli of these species on the physiological parameters of photobiont. Increased accumulation of Zn, Cd, Cu, and Ni in D. muscorum and of Zn and Ni in C. rei negatively affected contents of photosynthetic pigments, whereas concentrations of Pb had a positive effect in all lichen species. Moreover, pigment contents were positively associated with the concentrations of most examined elements in C. cariosa. The results indicate that even if chlorophyll contents reduced, its degradation does not progress. This suggests that metal stress may exert a negative effect on the synthesis rather than on the integrity of chlorophyll. Most importantly, lichen samples of each of the species from polluted sites proved to possess significantly higher FV/FM ratios than those from a reference site; moreover, the contents of elements of lichen thalli positively influenced this parameter. The efficient functioning of the algal component under heavy-metal stress conditions indicates that the examined lichens are well adapted to extremely contaminated substrates.

Certain lichens are effective colonisers of polluted sites. However, little is known about the tolerance of photobionts and the degree of mycobiont selectivity to photobionts relative to metal pollution. The present study recognises the genetic and anatomical diversity of Asterochloris photobionts in epigeic lichens, i.e. Cladonia cariosa, C. rei, and Diploschistes muscorum, in relation to a wide spectrum of soil pollution. In accordance with phylogenetic analysis, photobionts were clustered in 7 moderately- to well-supported clades, including 19 haplotypes. The mycobionts of all studied lichens demonstrated a low level of selectivity and were capable of associating with various Asterochloris lineages. This tendency was also expressed by the frequent (~25%) occurrence of multiple algal genotypes in a single thallus. This indicates that identified Asterochloris lineages are generally tolerant to heavy-metal pollution, and the low level of selectivity of mycobionts enables them to select the most suitable and/or available partner. The trend of increasing incidence of certain Asterochloris lineages and decreasing frequency of others along with increasing soil pollution was observed. This proves the superior adaptation of some photobionts to polluted sites. Such symbiotic plasticity constitute an adaptive feature necessary for the successful colonisation. High number of haplotypes at polluted sites could be the result of multiple introduction events from different areas during the initial stages of spontaneous succession. Regardless of the genetic pattern, Asterochloris cells were considerably smaller, and the density and compaction of cells in the algal layer were higher, in lichen specimens from polluted sites, indicating that photobiont characteristics may be closely dependent on heavy-metal pollution.

The epigeic lichens Cladonia rei and Diploschistes muscorum are effective heavy-metal-tolerant colonisers of highly polluted and disturbed sites. In this study we compare their bioaccumulation capacities, accumulation patterns, and responses to heavy-metal stress, as expressed in terms of cell membrane damage. We also aim at verifying the relationships between cell membrane damage and levels of soil pollution with heavy metals, and thereby to identify the bioindicative value of this physiological parameter. Total and intracellular concentrations of Zn, Pb, Cd, As, Cu, and Ni were measured in 140 samples of lichens and corresponding soil, collected from variously contaminated sites. Relative electrical conductivity (EC%) values were determined concurrently in the lichen samples. The studied lichens differ considerably in intracellular uptake susceptibility and the related reduction in membrane integrity. In C. rei thalli, more than half of Zn, Pb, Cd, and As loads are accumulated extracellularly, whereas D. muscorum exhibits a tendency towards intracellular accumulation of the same elements. This property is clearly reflected in cell membrane damage, which is considerably greater in the latter species irrespective of study site. This indicates that intracellular heavy-metal accumulation affects the level of cell membrane damage. Two soil pollution classes were distinguished for both lichens based on element contents in host-substrate samples. The losses of cell membrane integrity in lichen thalli are related to these classes. EC% values above 16 in C. rei and above 20 in D. muscorum suggest elevated levels of heavy metals in the soil. Consequently, this physiological parameter can serve as an early warning indicator for detection of elevated metal concentrations in soil. The biomonitoring method proposed here involves common and widespread lichen species and can be widely applied in post-industrial areas.

Lichens and bryophytes have commonly been used as bioindicators of environmental conditions, especially in relation to air quality. However, their diagnostic role in the assessment of soil pollution is relatively poorly recognised. The aim of this study was to find a pattern of cryptogamic biota structure associated with zinc and lead soil pollution and to thereby identify common signal species useful for bioindication purposes. The study area encompassed various types of anthropogenic and semi-natural habitats directly associated with the processing of Zn-Pb ores in southern Poland. Detailed analysis of cryptogamic biota with respect to the chemical parameters of the corresponding soil enabled us to identify three different pollution classes related to the concentration of heavy metals and four distinct groups of ecologically close species with similar responses to the prevailing level of pollution. The significant relationship between soil chemical parameters and cryptogamic biota structure implies the high bioindicative value of the defined lichen and bryophyte assemblages. Consequently, specific sets of distinct species reflecting levels of pollution were instrumental in the development of a practical tool. This approach may constitute a first step in soil quality assessment in a broad landscape scale. It provides an opportunity for preliminary verification of the sites that are potentially the most contaminated and which require further attention, for example, within the framework of restoration projects, reclamation interventions, or conservation strategies. The proposed bioindication approach involves common, widespread lichens and bryophytes, thus increasing the potential for its wide application in post-industrial areas associated with the mining and processing of Zn-Pb ores.

In this study, we examined oribatid fauna of strongly contaminated post-smelting dumps (southern Poland) that exist in the substrate and are associated with the most frequent lichen, Cladonia rei. Due to artificial origin of the substrate and the extremely high contamination with heavy metals, the studied dumps are unique in Europe in terms of unfavourable life conditions. In total, 2,936 specimens of Oribatida, representing 50 oribatid species, were sampled on 10 dumps and a reference site. Thalli of C. rei act as an island for soil oribatid mites on extremely contaminated post-smelting dumps. Both abundance and species richness of oribatid fauna collected from C. rei thalli were significantly greater than those recorded in the dump's substrate. The pool of oribatid species that was able to persist in extremely high doses of heavy metals was comparatively broad. However, only one species, Tectocepheus velatus, was able to achieve high abundances on all dumps. Three different responses of species (tolerant, sensitive, and indifferent) to heavy-metal contamination were recognised. Redundancy analysis indicated that highly increased levels of heavy metals, as well as K content, C/N ratio, and pH value, were the main factors that influence the composition and distribution of species. The concentrations of heavy metals (both essential elements (zinc) as well as xenobiotics (lead, cadmium) in T. velatus from the most contaminated dumps were not increased compared with those observed in moderately contaminated soils.

The paper presents the results of comprehensive analyses concerning the phenotypic variability of the primary thallus in five chosen Cladonia species in relation to habitat conditions. Morphology and anatomy, as well as the cortex ultrasculpture, of primary squamules were examined in individual specimens from sunny and shady populations of each species. The Mann-Whitney U test and Principal Component Analysis (PCA) revealed several clear correlations between the type of habitat and the organisation of squamules. Significant differences refer mainly to anatomical features, especially to epinecral layer frequency and cortex thickness. On the contrary, examination of the surface ultrasculpture of squamules under SEM did not show any important differences between populations. Regardless of the habitat factors, a full range of cortical surface rugosity in samples from both habitats was noticed, and a high variability often refers to a singular squamule. The results indicate that some features are largely modified by environmental factors and/or depend on the development stage of individual specimens, and examination of these relationships should be the first step towards the evaluation of the taxonomic usefulness of the considered features.

Lichens appear to be essential and effective colonisers of bare substrates including the extremely contaminated wastes of slag dumps. This study examines the metal accumulation capacity of epilithic lichens growing directly on the surface of artificial slag sinters. Four species representing different growth forms, i.e., crustose Candelariella aurella, Lecanora muralis, and Lecidea fuscoatra and fruticose Stereocaulon nanodes, were selected to evaluate the relationships between zinc, lead, cadmium, and nickel contents in their thalli and host substrates. Bioaccumulation factors of examined crustose lichens showed their propensity to hyperaccumulate heavy metals. Contrarily, concentrations of metals in fruticose thalli of S. nanodes were, as a rule, lower than in the corresponding substrates. This indicates that the growth form of thalli and degree of thallus adhesion to the substrate has a significant impact on metal concentrations in lichens colonising post-smelting wastes. Nonlinear regression models described by power functions show that at greater levels of Pb concentration in the substrate, the ability of C. aurella, L. muralis and L. fuscoatra to accumulate the metal experiences a relative decrease, whereas hyperbolic function describes a similar trend in relation to Ni content in S. nanodes. This phenomenon may be an important attribute of lichens that facilitates their colonisation of the surface of slag wastes. Electronic supplementary material The online version of this article (doi:10.1007/s00244-015-0180-5) contains supplementary material, which is available to authorized users.

This study aims to determine changes in the structure of cryptogamic vegetation of poor psammophilous grassland along a pollution gradient near a zinc smelter to evaluate the potential of species assemblages as bioindicators of soil condition. Lichens and bryophytes were examined in study plots along six transects in four distance zones, and the physicochemical properties of corresponding soil samples were analysed. Four different responses of species to substrate contamination were identified, with a distinct group of species resistant to and favoured by metal contamination. Although species richness decreases as one approaches the smelter, the gradual replacement of certain sensitive species by resistant ones was observed along the pollution gradient. The results enabled us to develop a useful tool to diagnose strongly polluted sites. Two different cryptogamic assemblages of well-recognised key species characteristic for strongly polluted and lightly polluted sites were distinguished. We conclude that cryptogamic community structure clearly corresponds to the degree of soil contamination, thus demonstrating high bioindicative value. The study confirmed the high relevance of the community approach in metal pollution biomonitoring.

OSYCZKA, P. 2013. A morphometric evaluation of the Cladonia chlorophaea group and allied taxa (Cladoniaceae, Ascomycota). — Herzogia 26: 49– 64. The Cladonia chlorophaea group represents a complex of morphologically and chemically similar taxa that have long perplexed taxonomists. Until now, few studies have systematically evaluated morphological characters in the group. A morphometric analysis of samples belonging to the taxa of the Cladonia chlorophaea and C. humilis groups as well as C. fimbriata, was performed using various fundamental statistical methods. It revealed four unambiguously distinct morphological aggregates of samples, corresponding to four well known, cosmopolitan species, namely C. asahinae, C. chlorophaea, C. fimbriata and C. humilis. The analysis highlights the most reliable diagnostic characters distinguishing the obtained groups which can be applied for species determination. A chemical race of C. asahinae with fumarprotocetraric acid chemosyndrome only is reported from Eu...

The phenomenon of mass occurrence of the lichen Cladonia rei in extremely contaminated post-smelting slag dumps was studied in relation to metal accumulation capacity of this lichen. The research was aimed to evaluate the relationships between element contents in the thalli and in the corresponding substrate. The study was conducted in terms of a wide spectrum of Zn, Cd, Pb and As contents. The concentrations of these elements in the lichen thalli and substrate samples were measured. Various regression models were considered to find the best fitted one that greatly reflects the dependencies. Various Cladonia species and the hyperaccumulator Diploschistes muscorum were also included in the study for comparison purposes. Specific non-linear regression models described by a power function reflected relationships between Zn and Cd contents in C. rei thalli and in the host substrate in the most reliable way. The relationship for As was also noted, but none significant model was found. Contrarily, Pb concentrations in the thalli varied independently of the metal levels in the substrate. Nevertheless, the concentrations of all measured heavy metals in C. rei thalli are relatively low considering the frequently enormous substrate contamination. Different Cladonia species demonstrated a generally similar accumulation capacity and could be considered as weak accumulators. The restrained accumulation pattern may be one of the important attributes of C. rei which facilitates its colonisation of extremely contaminated dumps. This finding highlights ecological importance of this species as stable and resistant pioneer in such affected sites.

This paper is the first report on Cladonia species from Ardahan, a north-eastern province of Turkey. A circumpolar boreal-low arctic species, Cladonia acuminata, rarely reported from Asia, and the recently described Cladonia monomorpha are reported as new for Turkey. Their detailed descriptions and taxonomical remarks are provided. Localities of other ascertained Cladonia species in the province supplement the knowledge of their distribution patterns in the country. In addition, the typically corticolous/lignicolous species Vulpicida pinastri is mentioned as also growing on primary squamules and podetia of C. pyxidata.

The study examines oribatid communities and heavy metal bioaccumulation in selected species associated with different microhabitats of a post-smelting dump, i.e. three lichen species of Cladonia with various growth forms and the slag substrate. The abundance of oribatids collected from the substrate was significantly lower than observed in lichen thalli. The morphology and chemical properties of lichens, and to some extent varying concentrations of heavy metals in thalli, are probably responsible for significant differences in oribatid communities inhabiting different Cladonia species. Some oribatids demonstrate the ability to accumulate zinc and cadmium with unusual efficiency, whereas lead is the most effectively regulated element by all species. A positive correlation was found between Zn content in all studied oribatids and their microhabitats. Oribatids exploring different food resources, i.e. fungivorous and non-fungivorous grazers, show considerable differences in bioconcentrations of certain elements. Electronic supplementary material The online version of this article (doi:10.1007/s11356-016-6100-z) contains supplementary material, which is available to authorized users.