Langfristige Auswirkungen ehemaliger Bewirtschaftungsvarianten auf die Diversit?t von Blütenpflanzen, Moosen und Flechten eines Kalkmagerrasens

Jeschke M., Kiehl K., Pfadenhauer J. and Gigon A. 2008. Long-term effects of former management on the diversity of vascular plants, mosses and lichens in a calcareous grassland. Bot. Helv. 118: 95 – 109. Long-term effects of grassland management on the species richness of vascular plants, mosses and lichens were studied in a calcareous grassland near Schaffhausen (Switzerland). Experimental plots were treated for 22 years (1977 – 1999) with different mowing regimes, annual burning in late winter, or abandonment. From 2000 onwards, all plots were mown annually in autumn. Five years later (2004), we studied the after-effects of the previous treatments by recording the cover of vascular plants, mosses and lichens in nested plots with sizes from 0.01 m² to 16 m². Vascular plant species richness as well as the number of species characteristic for calcareous grasslands were significantly lower in formerly abandoned or burnt plots than in regularly mown plots, with larger differences on small subplots. There was no significant effect of the previous mowing time (July vs. October) or mowing frequency (annual or every two years) on vascular plants. Mosses and lichens had been eliminated by annual burning between 1977 and 1999, and only few moss species re-colonized the formerly burnt plots until 2004. The species richness of mosses did not differ between formerly abandoned and mown plots, but there were more mesophytic and fewer xerophytic species in the formerly abandoned plots. Mesophytic mosses were also more abundant in plots that used to be mown every second year than in those mown annually. Our data indicate that even after five years of similar management, i.e. annual mowing in autumn, the previous long-term management had strong after-effects on species richness and species composition of calcareous grasslands. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Eingereicht am 19. M?rz 2008, Angenommen am 30. August 2008, Redaktion: Sabine Güsewell     

Lichens and mosses promote alternate stable plant communities in the New Jersey Pinelands

Severely burned areas in the pitch pine-dominated forests of the New Jersey Pinelands may remain open and only vegetated with mats of lichens and mosses and sparse, scattered vascular plants for many decades. We hypothesize that climate-driven alternation between moss and lichen domination of the cryptogam mats may foster and inhibit, respectively, vascular plant development. We propose that these processes are mediated by the inhibitory effects of lichens on seed germination and seedling establishment versus a facilitating effect of mosses. We tested the hypothesis by 1) detailed surveys of the composition of cryptogam mats and their association with vascular plants, 2) experimental studies of the effects of tissue leachates on seed germination, 3) observations of mycorrhizal infection in field-collected plants, and 4) experimental tests of seedling emergence from mats. Lichen dominance in the mats was correlated with low densities of vascular plants (graminoids and ericoid shrubs), thin organic horizons on the soil, and high levels of light availability; moss dominance was correlated with higher vascular plant densities, thicker organic horizons, and lower light. Tissue extracts of lichens strongly inhibited seed germination, while moss extracts had no effect. Similarly, mycorrhizal infection by both ecto- and endomycorrhizae was lower in plants growing within lichen mats than in moss mats or in bare soil. However, thick mats of both types of cryptogam inhibited seedling emergence. We observed that moss-dominated patches became overgrown with lichens during a series of very dry, hot summers during the study. These observations all support the hypothesis that fluctuating warm and dry versus cool and moist conditions allow alternative stable states (open cryptogam mats vs succession to pine forest) to develop.     

Effects of bryophytes and lichens on seedling emergence of alvar plants: evidence from greenhouse experiments

Emergence of seedlings of four alvar grassland species ( Arenaria serpyllifolia , Festuca ovina , Filipendula vulgaris and Veronica spicata ) in bryophyte and lichen carpets was analysed in a series of greenhouse experiments. The aspects investigated were: the influence of thickness of moss mats, both in dry and moist conditions, the effects of thick Cladonia spp. clumps, and of living vs dead moss shoots and lichen podetia. Overall, Festuca seedlings emerged best whereas the small-seeded species, Arenaria and Veronica , had the lowest emergence. Moisture had a significant effect only on the emergence of Festuca seedlings, which emerged better in the dry treatment than in the moist. A thick moss cover negatively affected seedling emergence of Arenaria and Veronica but did not affect the emergence of Festuca . Filipendula showed lower seedling emergence in both thick and thin moss than on bare soil only in the dry treatment, whereas in the moist treatment emergence did not differ among the three substrates. Arenaria seedlings emerged less in thick and thin moss than on bare soil in the dry treatment, whereas in the moist treatment emergence in the thin moss was not different from bare soil. Thus, in relatively dry environments even a thin moss cover may inhibit rather than facilitate seedling emergence. The lichen clumps inhibited only the emergence of the forbs. Both living moss shoots and lichen podetia inhibited emergence of Veronica seedlings but did not affect Festuca . In contrast, emergence in the presence of dead mosses and lichens did not differ from emergence in their absence for both species. Hence, inhibition of seedling emergence by bryophytes and lichens of at least some vascular plant species may be mediated by some biotic factor. However, the effect of differences in substrate properties on germination cannot be excluded     

Species richness of vascular plants, bryophytes and lichens in dry grasslands: The effects of environment, landscape structure and competition

We studied the relative importance of local habitat conditions and landscape structure for species richness of vascular plants, bryophytes and lichens in dry grasslands on the Baltic island of Öland (Sweden). In addition, we tested whether relationships between species richness and vegetation cover indicate that competition within and between the studied taxonomic groups is important. We recorded species numbers of vascular plants, bryophytes and lichens in 4 m² plots (n=452), distributed over dry grassland patches differing in size and degree of isolation. Structural and environmental data were collected for each plot. We tested effects of local environmental conditions, landscape structure and vegetation cover on species richness using generalized linear mixed models. Different environmental variables explained species richness of vascular plants, bryophytes and lichens. Environmental effects, particularly soil pH, were more important than landscape structure. Interaction effects of soil pH with other environmental variables were significant in vascular plants. Plot heterogeneity enhanced species richness. Size and degree of isolation of dry grassland patches significantly affected bryophyte and lichen species richness, but not that of vascular plants. We observed negative relationships between bryophyte and lichen species richness and the cover of vascular plants. To conclude, effects of single environmental variables on species richness depend both on the taxonomic group and on the combination of environmental factors on a whole. Dispersal limitation in bryophytes and lichens confined to dry grasslands may be more common than is often assumed. Our study further suggests that competition between vascular plants and cryptogams is rather asymmetric.     

Juniper Shade Enables Terricolous Lichens and Mosses to Maintain High Photochemical Efficiency in a Semiarid Temperate Sand Grassland

On a semiarid sand grassland (Festucetum vaginatae) colonised by juniper (Juniperus communis L.) shrubs terricolous lichens and mosses segregate strongly between microhabitats: certain species grow in the open grassland, others almost exclusively in the shade of junipers. The contrasting irradiances of these microhabitats influence much the metabolism of these organisms, and thus affect their small-scale distribution. This was confirmed by determining the efficiency of photochemical energy conversion by measuring chlorophyll a fluorescence parameters. In the open grassland maximum photochemical efficiency of photosystem 2 (PS2, F_v/F_m) declined from the humid spring to the hot and dry summer in all species, and this was caused by an increase in base fluorescence (F₀), but not by the decrease in fluorescence maximum (F_m). In summer, mosses and lichens growing in the open grassland generally possessed lower F_v/F_m than cryptogams growing in the shade cast by juniper shrubs. Thus mosses and lichens in the open grassland suffer lasting reduction in photochemical efficiency in summer, which is avoided in the shade of junipers. Juniper shrubs indeed influence the composition and small-scale spatial pattern of sympatric terricolous lichen and moss communities by—among others—providing a shelter against high light in summer.     

Spatial structure, rock type, and local environmental conditions drive moss and lichen distribution on calcareous boulders

Although mosses and lichens are a relevant component of the biota of rock habitats targeted for biodiversity conservation in Europe, the ecological factors driving their distribution are still poorly known. In this work, we examined the epilithic moss and lichen assemblages colonizing boulders of different types of calcareous rocks co-occurring in the same area in the Italian Alps. The goals were: (1) to evaluate if and to what extent different calcareous rocks host different assemblages; (2) to identify species associated to each rock type; (3) to quantify the relative importance of rock type, local environmental factors, and habitat spatial structure in explaining species distribution. Our results demonstrated that different calcareous rocks host different moss and lichen assemblages with some typical species, indicating that each rock type contributes to the total diversity of both mosses and lichens. Local environmental conditions influenced mosses and not lichens whose distribution is mainly associated to rock type. The patterns of both organism groups were also significantly related to habitat spatial structure, species assemblages tending to have a patchy distribution, which may reflect dispersal dynamics. Our results have implications for conservation: (1) each rock type may play a relevant role in maintaining the overall diversity contributing with unique assemblages and typical species; (2) the patchy distribution of both moss and lichen assemblages should warn from considering rock patches as a monotonous repetition of the same habitat across space.     

Differential effects of lichens,mosses and grasses on respiration and nitrogen mineralization in soils of the New Jersey Pinelands

In the New Jersey Pinelands, severely disturbed areas often do not undergo a rapid succession to forest; rather, a patchy cover of lichens, mosses and grasses persists for decades. We hypothesized that these plant covers affect soil microbial processes in different ways, and that these effects may alter the successional dynamics of the patches. We predicted that the moss and grass covers stimulate soil microbial activity, whereas lichens inhibit it, which may in turn inhibit succession. We collected soil cores from beneath each type of cover plus bare soil within two types of highly disturbed areas—sites subjected to hot wildfires, and areas mined for sand. Organic matter (OM) content, soil respiration and potential N mineralization were measured in the cores. Soils under mosses were similar to those under grasses; they accumulated more OM and produced more mineral N, predominantly in the form of ammonium, than either the bare soils or the soils beneath lichens. Mineralization under lichens, like that of the bare soils but unlike the soils beneath mosses or grasses, was dominated by net nitrification. These patterns were reproduced in experimentally transplanted moss and lichen mats. Mosses appear to create high-nutrient microsites via high rates of OM accumulation and production of ammonium, whereas lichens maintain low-nutrient patches similar to bare soil via low OM accumulation rates and production of mineral N predominantly in the mobile nitrate form. These differences in soil properties may explain the lack of vascular plant invasion in lichen mats, in contrast to the moss-dominated areas.     

Dynamics of cover, UV-protective pigments, and quantum yield in biological soil crust communities of an undisturbed Mojave Desert shrubland

Biological soil crusts are an integral part of dryland ecosystems. We monitored the cover of lichens and mosses, cyanobacterial biomass, concentrations of UV-protective pigments in both free-living and lichenized cyanobacteria, and quantum yield in the soil lichen species Collema in an undisturbed Mojave Desert shrubland. During our sampling time, the site received historically high and low levels of precipitation, whereas temperatures were close to normal. Lichen cover, dominated by Collema tenax and C. coccophorum, and moss cover, dominated by Syntrichia caninervis, responded to both increases and decreases in precipitation. This finding for Collema spp. at a hot Mojave Desert site is in contrast to a similar study conducted at a cool desert site on the Colorado Plateau in SE Utah, USA, where Collema spp. cover dropped in response to elevated temperatures, but did not respond to changes in rainfall. The concentrations of UV-protective pigments in free-living cyanobacteria at the Mojave Desert site were also strongly and positively related to rainfall received between sampling times (R² values ranged from 0.78 to 0.99). However, pigment levels in the lichenized cyanobacteria showed little correlation with rainfall. Quantum yield in Collema spp. was closely correlated with rainfall. Climate models in this region predict a 3.5–4.0 °C rise in temperature and a 15–20% decline in winter precipitation by 2099. Based on our data, this rise in temperature is unlikely to have a strong effect on the dominant species of the soil crusts. However, the predicted drop in precipitation will likely lead to a decrease in soil lichen and moss cover, and high stress or mortality in soil cyanobacteria as levels of UV-protective pigments decline. In addition, surface-disturbing activities (e.g., recreation, military activities, fire) are rapidly increasing in the Mojave Desert, and these disturbances quickly remove soil lichens and mosses. These stresses combined are likely to lead to shifts in species composition and the local extirpation of some lichen or moss species. As these organisms are critical components of nutrient cycling, soil fertility, and soil stability, such changes are likely to reverberate throughout these ecosystems.     

The influence of climatic legacies on the distribution of dryland biocrust communities

Predicting the distribution of biocrust species, mosses, lichens and liverworts associated with surface soils is difficult, but climatic legacies (changes in climate over the last 20 k years) can improve our prediction of the distribution of biocrust species. To provide empirical support for this hypothesis, we used a combination of network analyses and structural equation modelling to identify the role of climatic legacies in predicting the distribution of ecological clusters formed by species of mosses, lichens and liverworts using data from 282 large sites distributed across 0.6 million km² of eastern Australia. Two ecological clusters contained 87% of the 120 moss, lichen and liverwort species. Both clusters contained lichen, moss and liverwort species, but were dominated by different families. Sites where the air temperature increased the most over 20k years (positive temperature legacies) were associated with reductions in the relative abundance of species from the lichen (Peltulaceae and Teloschistaceae) and moss (Bryaceae) families (Cluster A species), greater groundstorey plant cover and lower soil pH. Sites where precipitation has increased over the past 20k years (positive precipitation legacy) were associated with increases in the relative abundance of lichen (Cladoniaceae, Lecideaceae and Thelotremataceae) and moss (Pottiaceae) families (Cluster B species) and lower levels of soil pH. Sites where temperatures have increased the most in the past 20k years suppressed the negative effects of plant cover on Cluster B by reducing plant cover. Increased intensity of grazing suppressed the negative effect of soil pH and the positive effect of soil carbon, on the relative abundance of Cluster B taxa. Finally, increasing temperature and precipitation legacies reduced the negative effect of soil pH on Cluster B. Understanding of the importance of climatic legacies improves our ability to predict how biocrust assemblies might respond to ongoing global environmental change associated with increasing land use intensification, increasing temperature and reduced rainfall.     

Unexpected side effects in biocrust after treating non‐native plants using carbon addition

Carbon addition has been proposed as an alternative to herbicide and manual removal methods to treat non‐native plants and reduce non‐target effects of treatments (e.g. impacts on native plants; surface disturbance). On Mojave Desert pavement and biocrust substrates after experimental soil disturbance and carbon addition (1,263 g C/m² as sucrose), we observed declines in lichens and moss cover in sucrose‐treated plots. To further explore this unforeseen potential side effect of using carbon addition as a non‐native plant treatment, we conducted biocrust surveys 5 and 7 years after treatments, sampled surface soils to observe if treatments additionally affected soil filamentous cyanobacteria, and conducted laboratory trials testing the effects of different levels of sucrose on cyanobacteria and desert mosses. Sucrose addition to biocrust plots reduced lichen and moss cover by 33–78% and species richness by 40–80%. Sucrose reduced biocrust cover in biocrust plots to levels similarly detected in pavement plots (<1%). While cyanobacteria in the field did not appear to be affected by sucrose, laboratory tests showed negative effects of sucrose on both cyanobacteria and mosses. Cyanobacteria declined by 41% 1 month after exposure to 5.4 g C/m² equivalent solutions. We detected injury to photosynthesis in mosses after 96 hour exposure to 79–316 g C/m² equivalent solutions. Caution is warranted when using carbon addition, at least in the form and concentration of sucrose, as a treatment for reducing non‐native plants on sites where conserving biocrust is a goal.     

Balancing positive and negative plant interactions: how mosses structure vascular plant communities

Our understanding of positive and negative plant interactions is primarily based on vascular plants, as is the prediction that facilitative effects dominate in harsh environments. It remains unclear whether this understanding is also applicable to moss–vascular plant interactions, which are likely to be influential in low-temperature environments with extensive moss ground cover such as boreal forest and arctic tundra. In a field experiment in high-arctic tundra, we investigated positive and negative impacts of the moss layer on vascular plants. Ramets of the shrub Salix polaris, herb Bistorta vivipara, grass Alopecurus borealis and rush Luzula confusa were transplanted into plots manipulated to contain bare soil, shallow moss (3 cm) and deep moss (6 cm) and harvested after three growing seasons. The moss layer had both positive and negative impacts upon vascular plant growth, the relative extent of which varied among vascular plant species. Deep moss cover reduced soil temperature and nitrogen availability, and this was reflected in reduced graminoid productivity. Shrub and herb biomass were greatest in shallow moss, where soil moisture also appeared to be highest. The relative importance of the mechanisms by which moss may influence vascular plants, through effects on soil temperature, moisture and nitrogen availability, was investigated in a phytotron growth experiment. Soil temperature, and not nutrient availability, determined Alopecurus growth, whereas Salix only responded to increased temperature if soil nitrogen was also increased. We propose a conceptual model showing the relative importance of positive and negative influences of the moss mat on vascular plants along a gradient of moss depth and illustrate species-specific outcomes. Our findings suggest that, through their strong influence on the soil environment, mat-forming mosses structure the composition of vascular plant communities. Thus, for plant interaction theory to be widely applicable to extreme environments such as the Arctic, growth forms other than vascular plants should be considered.     

Species richness correlations among primary producers in boreal forests

Close correlations in species numbers may make it possible to indirectly assess the species richness of difficult taxonomic groups by investigating indicator groups, for which data are more easily collected. We asked if species numbers correlate among the four dominating groups of primary producers in boreal forests (liverworts, macrolichens, mosses, and vascular plants) and if substrate affiliation of species (ground vs. other substrates), sample plot size (0.01–1000 m²), and stand age (young vs. old) influence correlation strength. We used three sets of study plots from northern Sweden each including wide ranges of species richness. Although there are large differences in the ecophysiology and substrate use of vascular plants and the two bryophyte groups (mosses and liverworts), we found strong positive correlations among them not previously reported from boreal forests. In contrast, no correlation in total species richness was found between macrolichens and the two bryophyte groups, despite large overlaps in their ecology. We suggest that the positive correlations among land plants (liverworts, mosses, and vascular plants) are linked to positive relationships between site moisture and species number for all three groups. In contrast, total species number of macrolichens has not been shown to be strongly associated with moisture. However, ground‐living lichens and mosses correlated negatively in old forests. This may relate to the inability of macrolichens to exploit shaded and wet old forest ground, a habitat that is used by many moss species. Furthermore, lichens and mosses of ‘other substrates’ correlated positively in old forests, probably because the amount of boulders was positively related to species richness in both groups. Generally, correlations became stronger with increasing plot size, whereas stand age had relatively little influence. We conclude that vascular plants could be used as an indicator group for species richness of mosses and liverworts in boreal landscapes.     

Comparisons of photosynthesis-related traits among understory lichens,mosses and vascular plant leaves in a high-elevation subalpine forest

对比高海拔亚高山森林林下地衣、藓类及维管植物叶片的光合相关性状 在高海拔亚高山森林中，许多地衣和藓类植物欣欣向荣，甚至成为林地表层的优势物种。尽管它们在森林生态系统中承担着不可替代的功能，但我们对其光合作用相关的功能性状水平知之甚少，对相同生境下藓类植物和维管植物的比较研究也较为匮乏。因此，本研究选择中国青藏高原东缘亚高山林下常见的3种地衣、3种藓类和4种维管植物，测定并对比了它们的碳、氮、磷含量及化学计量比、单位面积光合组织干重、叶绿素含量及光合光响应曲线的异同。研究结果表明，林下地衣的氮含量高于藓类植物，二者的氮、磷、叶绿素、光合速率及光合氮利用效率均低于维管植物叶片。不同种类地衣的光合相关性状与其生长环境相符，而林地表层的3种藓类植物和3种草本植物的功能性状在各自功能群中的种间差异均不显著。总之，这些结果表明高海拔亚高山林下地衣和藓类植物的优势与其光合相关性状关系不大，对严酷环境较强的适应能力可能是它们得以繁荣生长的关键。     

Response of feather moss associated N2 fixation and litter decomposition to variations in simulated rainfall intensity and frequency

Feather mosses in boreal forests form a dense ground‐cover that is an important driver of both nutrient and carbon cycling. While moss growth is highly sensitive to moisture availability, little is known about how moss effects on nutrient and carbon cycling are affected by the dynamics of moisture input to the ecosystem. We experimentally investigated how rainfall regimes affected ecosystem processes driven by the dominant boreal feather moss Pleurozium schreberi by manipulating total moisture amount, frequency of moisture addition and moss presence/absence. Moisture treatments represented the range of rainfall conditions that occur in Swedish boreal forests as well as shifts in rainfall expected through climate change. We found that nitrogen (N) fixation by cyanobacteria in feather mosses (the main biological N input to boreal forests) was strongly influenced by both moisture amount and frequency, and their interaction; increased frequency had greater effects when amounts were higher. Within a given moisture amount, N fixation varied up to seven‐fold depending on how that amount was distributed temporally. We also found that mosses promoted vascular litter decomposition rates, concentrations of litter nutrients, and active soil microbial biomass, and reduced N release into soil solution. These effects were usually strongest under low moisture amount and/or frequency, and revealed a buffering effect of mosses on the decomposer subsystem under moisture limitation. These results highlight that both the amount and temporal distribution of rainfall, determine the effect of feather mosses on ecosystem N input and the decomposer subsystem. They also emphasize the role of feather mosses in mediating moisture effects on decomposer processes. Finally, our results suggest that projected shifts in precipitation in the Swedish boreal forest through climate change will result in increased moss growth and N₂ fixation but a reduced dependency of the decomposer subsystem on feather moss cover for moisture retention.     

Host-specificity of moss-associated fungal communities in the Ny-Ålesund region (Svalbard,High Arctic) as revealed by amplicon pyrosequencing

Fungal communities play a significant role in regulating ecological processes in the Arctic tundra. However, the extent to which the Arctic moss species and host types (moss, lichen and vascular plant) determine the richness, diversity, and composition of fungal communities at a local scale has not been quantitatively explored. Using 454 pyrosequencing in the current study, we characterized the fungal communities associated with six moss species (Andreaea rupestris, Bryum pseudotriquetrum, Hymenoloma crispulum, Polytrichastrum alpinum, Racomitrium lanuginosum, and Sanionia uncinata) and compared them with fungal communities associated with lichens and vascular plants in the Ny-Ålesund region (High Arctic). Host-species preference had greater explanatory power than geographical factors (longitude, latitude, elevation) in shaping moss-associated fungal communities. Fungal communities associated with mosses differed significantly from those associated with vascular plants and lichens, suggesting specificity of the fungal communities among three host types. Pairwise comparison analysis also indicated that the relative abundance of many taxonomic groups (e.g., Chaetothyriales, Leotiales, Catenulifera, Alatospora, and Toxicocladosporium) significantly differed between mosses and the other two host types. These results suggest host factors significantly affect the distribution of the fungal species associated with these moss species in the local-scale Arctic tundra.     

Metabolic recovery of continental antarctic cryptogams after winter

The activation of metabolism after the winter period was investigated in several mosses and lichens in continental Antarctica. Thalli that were still in their over-wintering inactive state in early spring were sprayed artificially and the time-dependent activation of photosystem II (PSII), carbon fixation and respiration was determined using gas exchange and chlorophyll a fluorescence techniques. The investigated lichens recovered PSII activity almost completely within the first few minutes and gross photosynthesis was fully reactivated within a few hours. In contrast, photosynthesis took much longer to recover in mosses, which could indicate a general difference between the green-algal symbionts in lichens and moss chloroplasts. Only small and quickly reversible increased rates of respiration were observed for the foliose lichen Umbilicaria aprina from a more xeric habitat. In contrast, species occurring near persistent meltwater, such as the moss Bryum subrotundifolium and the lichen Physcia caesia, had highly increased respiration rates that were maintained for several days after activation. Calculation of the carbon balances indicated that the activation pattern strongly dictated the length of time before a carbon gain was achieved. It appears that the differences in recovery reflect the water relations of the main growth period in summer.     

Population size and fire intensity determine post‐fire abundance in grassland lichens

Questions: What is the variability in abundance of lichens on grassland soil between and within fields after prescribed fire? Is post‐fire lichen abundance an effect of pre‐fire population size? Location: Cedar Creek Natural History Area, Minnesota, USA. Methods: Lichen abundance, estimated as ground cover and dominated by Cladonia spp., was mapped in plots in two fields before prescribed burning on 06.10.2003 and 15.10.2003 for the first time since abandonment in the 1950s. The plots were resurveyed one year post‐fire. Results: Post‐fire cover of Cladonia spp. varied strongly between the fields, most likely due to different weather conditions between the burn events, which resulted in different fire intensities, one of low and one of high intensity. In the field that experienced the low intensity fire, post‐fire cover of Cladonia spp. was still relatively high, and showed a positive relationship with pre‐fire cover, while no such relationship was found after the high intensity fire. In that field Cladonia spp. experienced high mortality rates irrespective of pre‐fire cover. Conclusions: This study provides an example of how species response to disturbance can be a function of population size, but that this relationship can be non‐linear; lichens in grassland can survive a low intensity fire proportionally to pre‐fire population size, but experience high mortality rates above a fire intensity threshold. The applications of these results are that fire intensity matters to species response to prescribed fire, and that the persistence of climax lichen communities and biodiversity in the study system needs a broad range of fire intervals.     

Effects of reindeer on boreal forest floor vegetation: Does grazing cause vegetation state transitions?

Intensive reindeer grazing has been hypothesized to drive vegetation shifts in the arctic tundra from a low-productive lichen dominated state to a more productive moss dominated state. Although the more productive state can potentially host more herbivores, it may still be less suitable as winter grazing grounds for reindeer, if lichens, the most preferred winter forage, are less abundant. Therefore, such a shift towards mosses may have severe consequences for reindeer husbandry if ground-growing lichens have difficulties to recover. We tested if reindeer cause this type of vegetation state shifts in boreal forest floor vegetation, by comparing plant species composition and major soil processes inside and outside of more than 40-year-old exclosures. Lichen biomass was more than twice as high inside exclosures than in grazed controls and almost 5 times higher than in heavily grazed patches. Contrary to our predictions, net N mineralization and plant production were higher in the exclosures than in the grazed controls. The lack of response of phytometer plants in a common garden bioassay indicated that changed soil moisture may drive effects of reindeer on plant productivity in these dry Pine forest ecosystems.     

Determinants of Lichen Diversity in a Rain Forest Understory

Change in lichen diversity is often used as a bioindicator to estimate effects of atmospheric pollution, but natural variation in lichen cover and species richness can be very high. We examined the top-down effects of spore-consuming ants and the bottom-up effects of nutrient and light availability on lichen diversity associated with the leaf surface of the rain forest understory plant, Piper cenocladum. Plots containing P. cenocladum were randomly assigned to treatments in factorial experiments that included high and low light levels, nutrient enrichment, and presence and absence of the ant mutualist, Pheidole bicornis . At the conclusion of the experiments, plants were harvested and size of leaves, secondary metabolite content (amides), epiphyll cover, and the species richness of the lichens (which comprised 85% of the epiphyll community) were quantified. Epiphyll cover (mosses, liverworts, and lichens) was greater on plants that had ant-mutualists and balanced resources. Lichen species richness was greater for plants with balanced resources, particularly for those with high light availability. Relationships between toxins and lichen cover and richness were weak and unclear. In this system, natural sources of variation were reliable determinants of lichen diversity and both biotic and abiotic influences were important.     

Lichen secondary metabolites affect growth of <Emphasis Type="Italic">Physcomitrella patens</Emphasis> by allelopathy

Lichen secondary metabolites can function as allelochemicals and affect the development and growth of neighboring bryophytes, fungi, vascular plants, microorganisms, and even other lichens. Lichen overgrowth on bryophytes is frequently observed in nature even though mosses grow faster than lichens, but there is still little information on the interactions between lichens and bryophytes.In the present study, we used extracts from six lichen thalli containing secondary metabolites like usnic acid, protocetraric acid, atranorin, lecanoric acid, nortistic acid, and thamnolic acid. To observe the influence of these metabolites on bryophytes, the moss Physcomitrella patens was cultivated for 5 weeks under laboratory conditions and treated with lichen extracts. Toxicity of natural mixtures of secondary metabolites was tested at three selected doses (0.001, 0.01, and 0.1 %). When the mixture contained substantial amounts of usnic acid, we observed growth inhibition of protonemata and reduced development of gametophores. Significant differences in cell lengths and widths were also noticed. Furthermore, usnic acid had a strong effect on cell division in protonemata suggesting a strong impact on the early stages of bryophyte development by allelochemicals contained in the lichen secondary metabolites.Biological activities of lichen secondary metabolites were confirmed in several studies such as antiviral, antibacterial, antitumor, antiherbivore, antioxidant, antipyretic, and analgetic action or photoprotection. This work aimed to expand the knowledge on allelopathic effects on bryophyte growth.