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.

Across environmental gradients, community‐level functional traits of plants can change due to species turnover, intraspecific variation and their covariation. Studies on vascular plants suggest that species turnover is the main driver of trait variation across gradients, although intraspecific variation can also be important. However, there is limited knowledge about whether this holds for non‐vascular primary producers such as lichens and bryophytes. We hypothesized that intraspecific variation is more important for non‐vascular than for vascular primary producers because they lack specialized structures to maintain homeostasis and should therefore be more responsive to extrinsic factors. To assess the relative importance of species turnover versus intraspecific variation for vascular plants, lichens and bryophytes, we estimated species abundance and measured chemical (tissue nitrogen (N) and phosphorous (P) content, N:P ratio and pH) and non‐chemical (specific leaf or thallus area, dry matter content and water holding capacity) functional traits along an elevational gradient in alpine southern Norway. We calculated community‐weighted mean traits and quantified the relative contribution of species turnover, intraspecific variation and their covariation to total trait variation across the gradient. We found mixed support for our hypothesis: the contribution of intraspecific variation to total trait variation for N and N:P was higher in lichens than in vascular plants and bryophytes, but in general the contribution of intraspecific variation differed among functional traits and producer groups. Nutrient variables (N, P and N:P) were significantly impacted by intraspecific variation for vascular plants and lichens but not for bryophytes. Non‐chemical traits and pH were mainly driven by species turnover effects in all primary producer groups. Our results highlight that while nearly all studies on primary producer trait variation across environments have focused on vascular plants, trait variation of other largely neglected but ecologically important producer groups, such as lichens and bryophytes, may show very different responses to the same environmental factors. In order to fully understand how future environmental changes impact on community‐ and ecosystem‐level processes, traits of primary producers other than vascular plants—and their within‐species variation—need to be considered in systems where these groups are abundant. A free Plain Language Summary can be found within the Supporting Information of this article.

Summary1. There has been much recent interest in understanding how functional traits of vascular plant species drive ecological processes such as herbivory and litter decomposition. In plants, these two processes are often driven by the same or similar suites of traits and therefore correlate across species. However, few studies have considered how traits of plant-like life forms such as lichens determine species differences in their effects on ecological processes. This is despite the significant contribution of lichens to carbon and nutrient cycling in many environments. 2. We collected 28 lichen species that differed in their growth form, substrate type and capacity to fix N, and determined key traits for each species. For each species, we performed a feeding bioassay using the generalist snail Cepaea hortensis and carried out a laboratory bioassay to assess decomposability. We did these tests both with intact lichen material containing natural concentrations of carbon-based secondary compounds (CBSCs), and material that had been acetone rinsed to reduce concentrations of CBSCs, to evaluate the effect of CBSC on palatability and decomposability. 3. We found that reducing CBSC concentrations greatly increased palatability for 17 species, and decomposability of 10 species. However, decomposability was correlated with several lichen traits while palatability was not, regardless of whether or not CBSCs were removed, and we therefore found no relationship between decomposability and palatability across species. Decomposability and palatability both varied, but in contrasting directions, among N-fixing vs. non-fixing lichens, lichens with different growth forms and those from contrasting substrate types. As such, N-fixing lichens had higher decomposition rates but lower consumption rates than non-fixing lichens, while foliose species had higher decomposition rates but lower consumption rates than fruticose species. 4. Synthesis: We have shown that lichen CBSCs regulate key processes such as lichenivory and decomposition, that lichen decomposability but not palatability are related to traits, and that these two processes are unrelated across species. These results highlight the potential role of lichen species differences in influencing ecosystem processes relating to decomposition and nutrient cycling and the role that grazers may play in driving this.

THIS STUDY AIMS: (1) to quantify mollusc grazing on juvenile and mature thalli of the foliose epiphytic lichen Lobaria pulmonaria, and (2) to test the hypothesis inferring a herbivore defensive role of lichen depsidones in forests with indigenous populations of lichen-feeding molluscs. Lichens were transplanted in shaded and less shaded positions in each of two calcareous broadleaved deciduous forests, one poor in lichens, one with a rich Lobarion community. Preventing the access of molluscs significantly reduced the loss of juvenile L. pulmonaria, particularly in the naturally lichen-poor forest. Molluscs also severely grazed mature thalli in the lichen-poor forest, especially thalli placed under the more shading canopies. Furthermore, reducing the natural concentration of depsidones by pre-rinsing with acetone increased subsequent grazing significantly, showing that lichen depsidones function as herbivore defence in natural habitats. Our results suggest that mollusc grazing may play important roles in shaping the epiphytic vegetation in calcareous deciduous forests, and that recently established juvenile L. pulmonaria thalli seem to be particularly vulnerable.

Summary1. Species composition and biomass of epiphytic lichens varies along a complex environmental gradient from the ground to the top of the forest canopy. It is not known if this gradient, considered to be shaped by succession (age of bark surface) and climatic factors, is also influenced by invertebrate grazing. 2. To investigate the grazing hypothesis, the natural height ranges of four old-forest Lobaria species on tree trunks were quantified. These large foliose epiphytes with different secondary chemistry were transplanted onto Fraxinus excelsior trunks 0.5, 3 and 6 m above-ground in five broad-leaved deciduous forests in southern Norway. After 4.5 months of exposure to natural climbing gastropods, grazing was quantified. 3. Grazing pressure strongly increased with increasing proximity to the ground. At all heights, gastropods clearly preferred Lobaria scrobiculata followed by L. amplissima. Lobaria pulmonaria, highest in carbon-based secondary compounds (CBSCs), and L. virens, nearly deficient in CBSCs, were both much less grazed. Therefore, CBSCs cannot explain the preferences. According to existing literature the stictic acid complex, present in L. pulmonaria and L. scrobiculata, represents a herbivore defence when occurring in quantities as high as those in L. pulmonaria. The identity of the strong defence in the CBSC-deficient L. virens is unknown. 4. Gastropods' preference for these epiphytes mirrors the distribution of the lichens in nature. The highly palatable L. scrobiculata occurs mainly in localities with low gastropod abundance such as boreal forests and on bark with slightly lower pH. Lobaria amplissima occurs in gastropod-rich localities, but above the lower parts of the trunk. Lobaria pulmonaria and L. virens frequently grow down to a level of < 1 m above the ground. 5. Synthesis. Our results suggest that climbing gastropods play a role in determining the lower distribution limit of epiphytic lichens along a vertical canopy gradient and influence the spatial pattern of susceptible lichen species. By grazing lichens in a species-specific way, gastropods can shape epiphytic communities in broad-leaved deciduous forests.

Three epiphytic old forest lichens (Usnea longissima, Pseudocyphellaria crocata, and Lobaria pulmonaria) were transplanted along a natural shade-sun gradient comprising three successional stages in boreal spruce forests (dense young forest, open old forest, and clear-cut) for one summer. After harvest, extractable secondary compounds were analyzed by high-performance liquid chromatography, and the brown pigmentation in melanic species was quantified by reflectance measurements. Cortical compounds in all species increased from shady young forests to exposed clear-cuts. Usnic acid, the major cortical, secondary compound in U. longissima, showed consistently higher concentration in the clear-cut than in the two forested stands. Pseudocyphellaria crocata and L. pulmonaria, lacking extractable secondary compounds in the cortex, significantly increased their amounts of cortical melanins in well-lit stands. The medullary compounds showed more complex responses. Many were not influenced by environmental conditions during the transplantation, whereas the majority of those that responded showed the lowest concentration in clear-cut transplants. Only a few medullary compounds showed the highest concentration in the clear-cut, and at a low level of significance. The synthesis of UV-B-absorbing usnic acid and melanins seems to be part of an acclimation to increased light exposure. The medullary compounds in studied species barely function as solar screens despite their strong UV-B absorbance.

Summary1. There has been growing recent interest in the relative importance of within-species trait variation vs. across-species trait variation in vascular plants in determining total community-level trait variation across communities and environmental gradients. Recent studies on plant communities have generally found across-species variation to be more important than within-species variation, but comparable studies involving other functionally important biota, such as lichens, are largely lacking. 2. We used a fire-driven chronosequence involving 30 lake islands in northern Sweden to study how declining soil fertility during retrogression affects the functional traits of each of the dominant epiphytic lichen species growing on the trunks of the tree Betula pubescens. We measured several functional traits for the commonest lichen species on each island and used communityweighted measures to study the community-level responses of lichens to the gradient. 3. We found that as retrogression proceeds and soil fertility declines, thallus N and P concentrations and specific thallus mass (STM) increase, both within species and at the community level. Lichen secondary compounds showed contrasting within-species responses and were non-responsive at the whole community level. 4. By decomposing community-level measures of these traits across the gradient, we showed that for the three most responsive traits (N, P and STM), within-species variation was substantially more important than across-species variation. This emerges in part because lichen species composition was not very responsive to ecosystem retrogression, and because unlike vascular plants, lichens easily absorb elements over their entire surface, meaning that nutrient concentrations within lichen species are likely to more closely reflect nutrient availability. 5. We found that within-species variability drove the changes in community-weighted measures of lichen traits across a strong environmental gradient, which contrasts strongly with what we know from studies of vascular plants where across-species variation and species turnover is much more important. To understand how lichen functional traits at the community level respond to environmental factors, it is therefore essential to consider the responses of individual species, and the application of traits-based approaches to lichen communities needs to account for their considerable intraspecific variability.

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