Nitrogen deposition drives lichen community changes through differential species responses

Nitrogen (N) deposition has increased globally over the last 150 years and further increases are predicted. Epiphytic lichens decline in abundance and diversity in areas with high N loads, and the abundance of lichens decreases along gradients of increased deposition. Thus, although N is an essential nutrient for lichens, excessive loads may be detrimental for them. However, these gradients include many correlated pollutants and the mechanisms behind the decline are thus poorly known. The aim of this study was to assess effects of N deposition, alone, on the epiphytic lichen community composition in a naturally N‐poor boreal forest. For this purpose, whole spruce trees were fertilized daily with N at five levels, equivalent to 0.6, 6, 12.5, 25, and 50 kg N ha^?1 yr^?1, during four consecutive growing seasons (2006–2009), and changes in the abundance of lichens were monitored each autumn from the preceding year (2005). The studied lichen communities were highly dynamic and responded strongly to the environmental perturbation. N deposition detectably altered the direction of succession and reduced the species richness of the epiphytic lichen communities, even at the lowest fertilization application (6 kg N ha^?1 yr^?1). The simulated N deposition caused significant changes in the abundance of Alectoria sarmentosa, Bryoria spp., and Hypogymnia physodes, which all increased at low N loads and decreased at high loads, but with species‐specific optima. The rapid decline of A. sarmentosa may have been caused by the added nitrogen reducing the stability of the lichen thalli, possibly due to increases in the photobiont: mycobiont ratio or parasitic fungal attacks. We conclude that increases in nitrogen availability, per se, could be responsible for the reductions in lichen abundance and diversity observed along deposition gradients, and those community responses may be due to physiological responses of the individual species rather than changes in competitive interactions.     

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.     

Litterfall of epiphytic macrolichens in Nothofagus forests of northern Patagonia,Argentina: Relation to stand age and precipitation

Abstract: The objective of this study was to analyse how stand age and precipitation influence abundance and diversity of epiphytic macrolichens in southern beech Nothofagus forests, estimated by lichen litter sampling. Five sites of Nothofagus dombeyi (Mirbel) Oersted were selected in Nahuel Huapi National Park, Argentina. At each site, lichen fragments from the forest floor were collected at 12.5 m² plots in pairs of young and mature N. dombeyi forest. Additionally, two sites with multi‐aged subalpine Nothofagus pumilio (Poepp. et Endl.) Krasser forest were investigated in a similar manner. Average litterfall biomass per stand varied from less than 1 kg ha^?1 in a young low‐precipitation stand to a maximum of 20 kg ha^?1 in a mature high‐precipitation stand. In places with higher precipitation, litterfall biomass in N. dombeyi forest was considerably higher in old stands as compared with young ones. In places with less than 2000 mm of precipitation, differences in biomass were less pronounced. Old humid stands contained about twice as many taxa in the litter as old low‐precipitation stands and young stands in general. Mature stands in low‐precipitation sites only contained 17% of the litter biomass as compared with mature stands in high‐precipitation sites. Epiphytic lichen composition changed from predominating fruticose lichens (Usnea spp. and Protousnea spp.) in low‐precipitation stands to Pseudocyphellaria spp., Nephroma spp. and other foliose lichens, in the high‐precipitation stands. There were no clear differences in the proportion of fruticose and foliose lichens between young and old stands. Fruticose lichens dominated litter biomass in both N. pumilio sites.     

ERUMPENT CEPHALODIA,AN APPARENT CASE OF PHYCOBIAL INFLUENCE ON LICHEN MORPHOLOGY1,2,3

On the upper surface of some specimens of the lichen Lobaria cf. crosa (Eschw.) Nyl. there are shrub-like growths which have a morphology similar to that of a free-living fruticose lichen, Polychidium umhausense (Auersw.) Henss., and contain a cyanophyte phycobiont, Nostoc. A few growths also contain scattered colonies of a chlorophyte phycobiont, in which case the lichen tissue locally assumes the foliose form characteristic of the parent Lobaria thallus. The differentiation of dorsiventral lichen cortices and the formation of a lax medulla and distinct algal layer are correlated with the presence of the green phycobiont. The lichen substances atranorin, gyrophoric acid, and 4-0-methylgyrophoric acid occur in both, the foliose L. erosa thallus and the fruticose tissue. It is suggested that the fruticose structures are erumpent cephalodia which are derived from the outgrowth of internal, cephalodia and should not be considered, to be epiphytic colonies of the lichen Polychidium.     

Removal of secondary compounds increases invertebrate abundance in lichens

We investigated how lichen carbon-based secondary compounds (CBSCs) affect abundance of invertebrates in five lichen species growing on the forest floor (Cladonia rangiferina, Cladonia stellaris) or on tree trunks (Evernia prunastri, Hypogymnia physodes, Pseudevernia furfuracea). To do this, CBSCs were removed by rinsing lichen thalli in acetone (which has no adverse effects on the lichens) and the lichens were re-transplanted in their natural habitat. After 4 months there was higher abundance of mites, springtails and spiders in the three epiphytic lichens that had their CBSC concentrations reduced. The increase in predatory spiders following CBSC reduction suggests that the compounds have multitrophic consequences. The acetone treatment reduced the number of nematodes in four of the lichen species. Given that lichens serve as important habitats for a diverse range of invertebrates, increased knowledge of how lichen CBSCs may regulate their abundance helps us to better understand the role that lichens and their defence compounds play in structuring forest food webs.     

Lipid and Fatty Acid Composition of Tree-Growing and Terrestrial Lichens

A comparison of lipid and fatty acid composition was made of the tree-growing lichens Evernia prunastri (L.) Ach., Parmelia saxatilis (L.) Ach. and Hypogymnia physodes (L.) Ach. and the terrestrial species Cetraria islandica (L.) Ach. and Cladonia impexa Harm. In the terrestrial species the total lipid content varied strongly during spring, while the tree-growing species showed much less variation. Phospholipid and sterol content of all lichens was unusually low. Monoglycosyl diglyceride was absent from Parmelia saxatilis. Fatty acids common to higher plants as palmitic, stearic, oleic, linoleic and linolenic acid were invariably present in all lichen species. In addition large quantities of extra-long chain fatty acids like behenic acid, eicosadienoic acid and cyclic aliphatic lichen acids were present in the terrestrial species. The degree of (poly) unsaturation decreased in the order Evernia prunastri, Parmelia saxatilis. Fatty acids common to higher plants as palmitic, stearic, impexa, which decrease was compensated by an increase in extralong chain fatty acids and lichen acids. It is suggested that the lichen acids are of adaptive value for lichen species growing in the terrestrial habitats, which were characterized by extreme diurnal temperature variations. Just as the polyunsaturated fatty acids, lichen acids guarantee at lower temperatures a high flexibility of the membranes involved, at the same time as they are less susceptible to photo-oxidation at the high daytime temperatures.     

Light use efficiency of dry matter gain in five macro-lichens: relative impact of microclimate conditions and species-specific traits

Relations between irradiance (I) and lichen growth were investigated for five macro‐lichens growing at two sites in Sweden. The lichens represented different mycobiont–photobiont associations, two morphologies (foliose, fruticose) and two life forms (epiphytic, terricolous). The lichens were transplanted at two geographically distant sites in Sweden (1000 km apart) from Sept 1995 to Sept 1996 in their typical microhabitats, where microclimate and growth were followed. Between April/May and Sept 96, the terricolous species had a dry matter gain of 0·2 to 0·4 g (g DW)^–1 and the epiphytes 0·01 to 0·02 g (g DW)^–1. When related to area, growth amounted to 30 to 70 g m^?2 for the terricolous species and to 1 to 4 g m^?2 for the epiphytes. There was a strong correlation between growth and intercepted irradiance when the lichens were wet (I_wet), with 0·2 to 1·1 g lichen dry matter being produced per MJ solar energy. Across the 10 sets of transplants, light use efficiencies of dry matter yield (e) ranged between 0·5 and 2%, using an energy equivalent of 17·5 kJ g^?1 of lichen dry matter. The higher productivity of the terricolous species was due to longer periods with thallus water contents sufficient for metabolic activity and because of the higher mean photon flux densities of their microhabitat. A four‐fold difference in photosynthetic capacity among the species was also important. It is concluded that lichen dry matter gain was primarily related to net carbon gain during metabolically active periods, which was determined by light duration, photon flux density and photosynthetic capacity.     

A Fourier Transform-Raman Spectroscopic Study of Lichen Strategies on Granite Monuments

The biodeterioration of granite by Lecidea fuscoatra (L.) Ach., Porpidia cinereoatra (Ach.) Hertel & Knoph, and P. macrocarpa (DC.) Hertel & Schwab growing in the same environmental conditions has been studied by using Fourier transform-Raman spectroscopy. Results were significantly different for the three species, with P. cinereoatra being the most active biodeteriorative lichen. This lichen was also the only one in which calcium oxalate and gypsum were identified spectroscopically. Physical disturbances to the substratum were evidenced in all lichens studied by the incoporation of material, such as quartz and feldspar, into their thalli, but this phenomenon varied considerably from species to species. The results indicate that lichen species can adopt different strategies and can have different biodeteriorative effects on granite, independently of environmental conditions and substrate.     

Differential effects of reindeer on high Arctic lichens

Abstract. We studied the effects of Svalbard reindeer on the abundance of lichens in Spitsbergen. A survey was carried out in 14 areas with contrasting reindeer densities. Separate cover estimates for crustose, fructose and foliose lichens were taken in each area, and related to the density of reindeer pellet groups, a measure of reindeer density. Dominant macro lichen families were identified in 10 areas, and a full record of macrolichen species was taken in four additional areas. Variation in reindeer density is partially due to past overhunting, and subsequent incomplete recovery, releasing some areas from reindeer grazing for 100–200 yr. The cover of fruticose lichens was negatively related to reindeer pellet group density, indicating suppression by Svalbard reindeer. This makes their impact comparable to other members of the Rangifer genus around the northern hemisphere. The generally recorded low abundance of lichens in the diet of Svalbard reindeer compared to other Rangifer species, therefore, was interpreted as the depletion of fruticose lichens in Spitsbergen, and a subsequent switch to alternative foods. Of all fruticose lichens, Stereocaulon spp. appeared least sensitive to grazing. Crustose and foliose lichen cover was independent of reindeer pellet group density. The cover of crustose lichens was significantly related to latitude, with greater cover in more northern areas. Foliose lichens were more abundant in places where moss cover was high. We conclude that the impact of Svalbard reindeer on lichens is dependent on growth form, with fruticose lichens suffering from grazing, whereas foliose lichens might indirectly benefit from higher densities of reindeer or, like crustose lichens, be controlled by other factors.     

Modelling hydration and photosystem II activation in relation to in situ rain and humidity patterns: a tool to compare performance of rare and generalist epiphytic lichens

A dynamic water and activity model was developed to assess how efficiently lichens can exploit in situ rain and humid air. The capacity to rehydrate and activate photosynthesis [i.e. photosystem II (PSII)] by these water sources was compared among four hydrophilic and one generalist epiphytic lichen. Hydration status, potential (instant activation) and realized (delayed activation) day‐light activity were simulated using a model based on species‐specific hydration, PSII activation characteristics and in situ water content for Platismatia norvegica in three microclimatic scenarios. The results showed that delayed PSII activation could have profound effects on lichens' ability to exploit environmental water sources. During rain, realized activity was reduced by 19, 34 and 56% compared to simulations assuming instant activation for three hydrophilic lichens in the driest microclimate. During humid air, the reduction was 81% for the most extreme species and scenario, because of slow hydration and low equilibrium water content. Many and brief hydration events may thus hamper species with slow activation and fast desiccation kinetics. No evidence of compensation by a ‘water‐holding’ morphology was observed among studied species. The developed model may provide a tool for identifying suitable habitats for long‐term persistence of lichens with physiological constraints.     

Tree and stand level variables influencing diversity of lichens on temperate broad-leaved trees in boreo-nemoral floodplain forests

Tree and stand level variables affecting the species richness, cover and composition of epiphytic lichens on temperate broad-leaved trees (Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra, and U. laevis) were analysed in floodplain forest stands in Estonia. The effect of tree species, substrate characteristics, and stand and regional variables were tested by partial canonical correspondence analysis (pCCA) and by general linear mixed models (GLMM). The most pronounced factors affecting the species richness, cover and composition of epiphytic lichens are acidity of tree bark, bryophyte cover and circumference of tree stems. Stand level characteristics have less effects on the species richness of epiphytic lichens, however, lichen cover and composition was influenced by stand age and light availability. The boreo-nemoral floodplain forests represent valuable habitats for epiphytic lichens. As substrate-related factors influence the species diversity of lichens on temperate broad-leaved trees differently, it is important to consider the effect of each tree species in biodiversity and conservation studies of lichens. Nomenclature Randlane et al. (2007) for lichens; Leht (2007) for vascular plants.     

FREQUENCY,DIVERSITY AND ECOLOGICAL STRATEGIES OF EPIPHYTIC LICHENS IN THE SWISS CENTRAL PLATEAU AND THE PRE-ALPS

To identify representative quantitative criteria for the creation of a future Red List of epiphytic lichens, 849 trees in 132 long-term ecological observation plots in the Swiss Central Plateau and the Pre-Alps were surveyed by standard sampling. Based on the trees, frequency data of the lichen taxa observed are described by the log series model, indicating the controlling effect of few ecological factors. Based on the plots, four classes of scarcity, each comprising 25% of the species, were established. As a contribution to the development of a national, representative survey of lichens, α-diversity (species richness, species density) andβ-diversity (dissimilarity) were calculated in terms of region, vegetation formation, vegetation belt and for their combinations. Differences in lichen diversity between the Central Plateau and the Pre-Alps were caused by the bigger elevational range in the Pre-Alps, which resulted in a higher species richness. α-Diversity of forest and non-forest were similar, whereas each vegetation formation showed one third of its species restricted to it. The contributions to the total lichen diversity of crustose, foliose and fruticose as well as of generative and vegetative species was calculated. Specific features along the altitudinal gradient of vegetation belts emerged: the percentage of crustose and generative lichens declined with every altitudinal step, increased in fruticose and vegetative lichens, and was the same in foliose species.     

哀牢山山地森林不同附生地衣功能群的水分关系和光合生理特征

附生地衣是热带和亚热带山地森林生态系统中重要的结构性组分, 在生物多样性保护、环境监测、养分循环中发挥着重要作用。附生地衣按共生藻、生活型和繁殖策略的不同可划分为不同的功能群, 不同附生地衣功能群的分布格局存在较大的差异, 然而其生理生态机制仍不清楚。该研究以我国西南地区哀牢山亚热带山地森林中的附生地衣优势类群为研究对象, 对该地区蓝藻地衣、阔叶地衣、狭叶地衣及枝状地衣4种功能群的8种附生地衣的水分关系、光合生理特征等进行了测定分析, 结果显示: 不同功能群附生地衣的持水力和失水速率均存在差异, 其中蓝藻地衣具有较高的最大水分含量, 而枝状地衣的失水速率较快; 过高和过低的水分含量都会抑制附生地衣的光合作用, 但抑制程度有所差异; 蓝藻地衣的光合作用最适水分含量比较高, 表明它们的光合生理活动对水分条件要求较高, 所以它们偏好潮湿的生境, 同时蓝藻地衣的光补偿点比较低但光饱和点却不低, 反映出它们具有较宽的光强适应范围, 所以蓝藻地衣能够同时分布于强光和弱光生境中; 枝状地衣的光合最适水分含量较低, 表明它们的光合生理活动对水分条件要求不是很高, 能够适应较为干旱的环境, 同时枝状地衣的光补偿点和光饱和点都很高, 说明它们的光合生理活动对光照条件要求比较高, 所以它们广泛分布于强光生境中; 阔叶和狭叶地衣的光补偿点比较高, 说明它们更适应有充足光照条件的生境。     

Predicting growth of mat‐forming lichens on a landscape scale – comparing models with different complexities

During the 20th century, forestry practices has adversely affected lichen‐rich habitats. Mat‐forming lichens are important components of the vegetation of boreal and arctic ecosystems and are the main reindeer forage during the winter. To support the long‐term management of lichens in such habitats we developed models for predicting the growth of two common species. The lichens were transplanted across northern Scandinavia along a west‐east gradient varying in precipitation, temperature and irradiance. Growth was recorded seasonally over 16 months and ranged from ?4.8 to 34.6% and ?12.7 to 34.7% dry weight change for Cetraria stellaris and Cladina islandica, respectively. Growth was light limited below canopies with more than ca 60% cover and highest at the more humid sites when light levels were optimal. The models were based on various meteorological parameters, irradiance, physiological data and lichen hydration status; the latter was derived from a recently developed lichen hydration model. Our models' abilities to predict growth, both annually and seasonally (i.e. in summer), were evaluated in relation to their complexity and their potential usefulness from a management perspective. One parameter related to irradiance (the logarithm of site openness) was valuable in the prediction of annual growth for both species and could, in combination with precipitation, explain 52% of the variation in annual growth for C. stellaris and, in combination with total wet time and the irradiance received while wet, explain 66% of the variation in annual growth for C. islandica. The best simplified model explained 43% of the variation in annual growth for C. stellaris, using stem basal area and the annual normal temperature, and 24% for C. islandica using basal area alone. It is concluded that ensuring sufficient irradiance below the forest canopy is of crucial importance in the long‐term management of mat‐forming lichens and that simplified models can be used to identify appropriate habitats.     

Water relations and photosynthetic characteristics in different functional groups of epiphytic lichens in montane forest of Ailaoshan

Aims There are abundant epiphytic lichens in the tropical and subtropical montane forest ecosystems, which are important components of forest canopy and play a vital role in biodiversity conservation, environmental monitoring and nutrient cycling. In accordance with photobiont type, growth form and reproductive strategy, the epiphytic lichens can be divided into different functional groups, with different distribution patterns. In this study we aim to explain this phenomenon from the perspective of physiological ecology. Methods The maximum water content, water loss curves, photosynthetic water and light response curves were determined in four epiphytic lichen functional groups, including cyanolichens, fruticose lichens, broadly lobed foliose lichens and narrowly lobed foliose lichens. Important findings The functional characteristics of epiphytic lichens influence their maximum water-holding capacity and rate of water loss. The cyanolichens have higher maximum water content, while the fruticose lichens have a faster water loss. The cyanolichens that are widely distributed in the moist habitats require particularly high moisture for their photosynthetic activities; their optimal water content for photosynthesis is higher in comparison with other groups. They also have a low light compensation point and a high light saturation point, which explain the wide range of light intensity of the habitat. The fruticose lichens, widely distributed in the relatively arid habitats with high irradiance, have high light compensation point and light saturation point, and low optimum water content for photosynthesis. The broadly lobed foliose lichens and the narrowly lobed foliose lichens have a high light compensation point and light saturation point; they preferably occur in habitats with strong light.     

Estimating plant abundance using inflated beta distributions: Applied learnings from a lichen–caribou ecosystem

Quantifying abundance and distribution of plant species can be difficult because data are often inflated with zero values due to rarity or absence from many ecosystems. Terrestrial fruticose lichens (Cladonia and Cetraria spp.) occupy a narrow ecological niche and have been linked to the diets of declining caribou and reindeer populations (Rangifer tarandus) across their global distribution, and conditions related to their abundance and distribution are not well understood. We attempted to measure effects related to the occupancy and abundance of terrestrial fruticose lichens by sampling and simultaneously modeling two discrete conditions: absence and abundance. We sampled the proportion cover of terrestrial lichens at 438 vegetation plots, including 98 plots having zero lichens. A zero‐inflated beta regression model was employed to simultaneously estimate both the absence and the proportion cover of terrestrial fruticose lichens using fine resolution satellite imagery and light detection and ranging (LiDAR) derived covariates. The probability of lichen absence significantly increased with shallower groundwater, taller vegetation, and increased Sphagnum moss cover. Vegetation productivity, Sphagnum moss cover, and seasonal changes in photosynthetic capacity were negatively related to the abundances of terrestrial lichens. Inflated beta regression reliably estimated the abundance of terrestrial lichens (R² = .74) which was interpolated on a map at fine resolution across a caribou range to support ecological conservation and reclamation. Results demonstrate that sampling for and simultaneously estimating both occupancy and abundance offer a powerful approach to improve statistical estimation and expand ecological inference in an applied setting. Learnings are broadly applicable to studying species that are rare, occupy narrow niches, or where the response variable is a proportion value containing zero or one, which is typical of vegetation cover data.     

新疆博格达山岩面生地衣群落结构特征

根据对新疆博格达山岩面生地衣群落20个样点(20m×20m)调查的数据,以各地衣种的盖度为指标结合双向指示种分析方法(TWINSPAN)和除趋势对应分析法(DCA)对博格达山岩面生地衣群落进行数量分类并分析了群落结构特征及其多样性和相似性。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研究。结果表明,TWINSPAN分析和DCA排序将分布在博格达山的37种岩面生地衣分为以下5个群丛。群丛1:斑纹网衣(Lecidea tessellate Florke)+粉芽盾衣(Peltula euploca(Ach.)Poelt)+杜瑞氏黄梅(Xanthoparmelia durietzii Hale)群丛,有25个种,总覆盖度为30.145%,多样性为4.025;群丛2:袋衣(Hypogymnia physodes(L.)Nyl.)+白边平茶渍(Aspicilia sublaqueata(H.Magn.)J.C.Wei)+砖孢胶衣(Collema subconveniens Nyl.)群丛,有17个种,地衣总盖度为15.885%,多样性为3.196;群丛3:聚茶渍(Lecanora accumulate H.Magn.)+丽石黄衣(Xanthoria elegans(Link)Th.Fr.)+亚洲平茶渍(Aspicilia asiatica(H.Magn.)Yoshim.)群丛,有30个种,地衣总盖度为37.87%,多样性为4.357;群丛4:中华石果衣(Endocarpon sinense H.Magn.)+伴藓大孢蜈蚣衣(Physconia muscigena(Ach.)Poelt.)+垫脐鳞衣(Rhizoplaca melanophthalma(DC.)LeuckertPoelt)群丛,有24个种,地衣总盖度为30.458%,多样性为3.912;群丛5:石胶衣(Collema flaccidum(Ach.)Ach.)+短绒皮果衣(Dermatocarpon vellereum Zschacke)+绿黑地图衣(Rhizocarpon viridiatrum(Wulfen)Korber.)群丛,有18个种,地衣总盖度为19.331%,多样性为3.515。CCA排序结果反映,该地区岩面生地衣的分布与海拔高度、光照强度、岩石pH和人为干扰有关,其中影响最大的因素是海拔高度,其次为光照强度和干扰。坡向和岩石大小对地衣种类分布的影响不显著。     

Water vapor uptake and photosynthesis of lichens: performance differences in species with green and blue-green algae as phycobionts

Summary Dry lichen thalli were enclosed in gas exchange chambers and treated with an air stream of high relative humidity (96.5 to near 100%) until water potential equilibrium was reached with the surrounding air (i.e., no further increase of weight through water vapor uptake). They were then sprayed with liquid water. The treatment took place in the dark and was interrupted by short periods of light. CO₂ exchange during light and dark respiration was monitored continuously. With no exception water uptake in all of the lichen species with green algae as phycobionts lead to reactivation of the photosynthetic metabolism. Further-more, high rates of CO₂ assimilation were attained without the application of liquid water. To date 73 species with different types of Chlorophyceae phycobionts have been tested in this and other studies. In contrast, hydration through high air humidity alone failed to stimulate positive net photosynthesis in any of the lichens with blue-green algae (Cyanobacteria). These required liquid water for CO₂ assimilation. So far 33 species have been investigated, and all have behaved similarly. These have included gelatinous as well as heteromerous species, most with Nostoc phycobionts but in addition some with three other Cyanophyceae phycobionts. The same phycobiont performance differences existed even within the same genus (e.g. Lobaria, Peltigera) between species pairs containing green or blue-green phycobionts respectively. Free living algae also seem to behave in a similar manner. Carbon isotope ratios of the lichen thalli suggest that a definite ecological difference exists in water status-dependent photosynthesis of species with green and blue-green phycobionts. The underlying biochemical or biophysical mechanisms are not yet understood. Apparently, a fundamental difference in the structure of the two groups of algae is involved.     

Lichen diversity and red-listed lichen species relationships with tree species and diameter in wooded meadows

This study provides a unique large dataset of total epiphytic lichen diversity (fruticose, foliose and crustose species) and composition on 1,294 trees of 17 tree species in wooded meadows in Sweden and Estonia, the Baltic region. The inventory (25,380 observations and 246 lichen taxa) clearly illustrated that Ulmus minor, Quercus robur and Fraxinus excelsior contributed most significantly to epiphytic lichen richness and number of red-listed species. In Sweden, average single tree α richness was 22.2 on Ulmus (only in Sweden), 21.6 on Quercus (25.0 in Estonia) and 19.8 on Fraxinus (16.7 in Estonia), respectively. Ulmus hosted on average one red-listed species per tree, compared with 0.7 on Fraxinus (0.6 in Estonia), 0.4 on Quercus (0.7 in Estonia) and only 0.05 on Betula (same in Estonia). Lichen species composition and the average number of red-listed lichens were influenced by tree diameter on Fraxinus and Quercus, whilst no such pattern was evident on Ulmus. Randomized species accumulation curves of the dominating tree species illustrated that Fraxinus, Quercus and Ulmus supported α dominated lichen communities where individual trees hosted a substantial part of the total richness. Betula, on the other hand, supported β dominated communities where individual trees tended to be dissimilar and, therefore, more of the total richness existed as species turnover among host trees. Lichen species composition was influenced by tree species, and most notably, lichen species on Ulmus had a strong consistent clumping in ordination graphs, with many rare and red-listed lichens. The broadleaved deciduous trees within the wooded meadows clearly contribute greatly to the biodiversity of the Baltic region.     

Communities Adjust their Temperature Optima by Shifting Producer-to-Consumer Ratio, Shown in Lichens as Models: II. Experimental Verification

The community adaptation hypothesis [7] predicts that lichens, simple communities of microorganisms, can adapt to a wide range of thermal regimes by regulating the ratio of primary producers (algae) and consumers (fungi): R_p/c. To test this hypothesis, we determined R_p/c values by image analysis of cross sections of herbarium specimens of the lichen Cladina rangiferina, which is widely distributed between the Arctic and the tropics. We found that R_p/c for C. rangiferina increases with summer temperature by more than one order of magnitude, consistent with the hypothesis. To assess the ecological significance of community adaptation (R_p/c regulation), other adaptive mechanisms (e.g., photobiont substitution, genetic adaptation, and photosynthetic acclimation in North American Cladina spp.) were studied. Laboratory investigations with algae and fungi isolated in culture from live specimens suggested that the role of these mechanisms is relatively minor and cannot account for the high degree of lichen adaptability.