The weight of the crust: Biomass of crustose lichens in tropical dry forest represents more than half of foliar biomass

In recent years, our ecological knowledge of tropical dry forests has increased dramatically. However, the functional contributions of whole ecosystem components, such as lichens, remain mostly unknown. In these forests, the abundance of epiphyte crustose lichens is responsible for the characteristic white bark on most woody plants, conspicuous during the dry season, but the amount of resources that the lichen component represents remains unexplored. We estimated lichen biomass in a Mexican tropical dry forest using the bark area of trees, the dry mass of lichens per unit area and the percentage of bark covered by lichens, together with previously known tree densities. The lowest 2.5 m of the forests main trunks contained 188 kg/ha of lichen biomass, with lichens covering 85% of the available bark for trees <12 cm DBH and 38% for trees >12 cm. Total epiphytic lichen biomass was 1.34–1.99 Mg/ha. Lichen biomass represented 61% of the foliar biomass in the forest. To our knowledge, this is the first time that a lichen biomass estimate is provided for an ecosystem in which crustose lichens are the dominant lichen growth form. Crustose lichens are typically considered to contribute little to the total lichen biomass and to be difficult to include in ecological analyses. The high lichen biomass in this ecosystem implies a significant ecological role which so far is unexplored. We suggest the crustose lichen component should not be underestimated a priori in ecological studies, especially in ecosystems with abundant lichen cover.     

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.     

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.     

Growth of foliose lichens: a review

This review considers various aspects of the growth of foliose lichens including early growth and development, variation in radial growth rate (RaGR) of different species, growth to maturity, lobe growth variation, senescence and fragmentation, growth models, the influence of environmental variables, and the maintenance of thallus symmetry. The data suggest that a foliose lichen thallus is essentially a ‘colony’ in which the individual lobes exhibit a considerable degree of autonomy in their growth processes. During development, recognisable juvenile thalli are usually formed by 15 months to 4 years while most mature thalli exhibit RaGR between 1 and 5 mm yr⁻¹. RaGR within a species is highly variable. The growth rate-size curve of a foliose lichen thallus may result from growth processes that take place at the tips of individual lobes together with size-related changes in the intensity of competition for space between the marginal lobes. Radial growth and growth in mass is influenced by climatic and microclimatic factors and also by substratum factors such as rock and bark texture, chemistry, and nutrient enrichment. Possible future research topics include: (1) measuring fast growing foliose species through life, (2) the three dimensional changes that occur during lobe growth, (3) the cellular changes that occur during regeneration, growth, and division of lobes, and (4) the distribution and allocation of the major lichen carbohydrates within lobes.     

A bird’s eye view of the peppered moth

Industrial melanism in Biston betularia is one of the best known examples of the role of natural selection in evolution and has received considerable scrutiny for many years. The rise in frequency of the dark form of the moth (carbonaria) and a decrease in the pale form (typica) was the result of differential predation by birds, the melanic form being more cyptic than typica in industrial areas where the tree bark was darkened by air pollution. One important aspect of early work evaluating the relative crypsis of the forms of B. betularia on tree trunks with different lichen flora was the reliance on human observers. Humans, however, do not have the same visual capabilities as birds. Birds have well‐developed ultraviolet (UV) vision, an important component of their colour processing system that affects many aspects of behaviour, including prey detection. We examined the UV characteristics of the two forms of B. betularia and a number of foliose and crustose lichens. In human visible light the speckled form typica appeared cyptic when seen against a background of foliose lichen, whereas the dark form carbonaria was conspicuous. Under UV light the situation was reversed. The foliose lichens absorbed UV and appeared dark as did carbonaria. Typica, however, reflected UV and was conspicuous. Against crustose lichens, typica was less visible than carbonaria in both visible and UV light. These findings are considered in relation to the distribution and recolonization of trees by lichens and the resting behaviour of B. betularia.     

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.     

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.     

Bacterial ecology of an old-growth douglas fir canopy

Microbial populations associated with the major substrates of the canopy of a single 70 m old-growth Douglas fir were studied to determine potential activities. Seasonal samples from bark, foliage, epiphytic moss, lichens, and litter accumulations were collected to: (a) obtain population data, (b) isolate the major groups of microorganisms present, (c) measure enzymatic activities associated with cellulose and xylan degradation, and (d) examine the potential for nitrogen fixation. We tested 562 bacterial isolates for utilization of 25 compounds associated with the canopy substrates, and for activities in nitrogen and sulfur cycle transformations. Total bacterial populations, reflecting seasonal temperature and moisture conditions, were lowest on bark and foliage [21–266×10³ colony-forming units (CFU/g)] and highest on moss and lodged litter (19–610×10⁵ CFU/g). Lichens contained intermediate numbers of bacteria (3.3–270×10⁵ CFU/g). The majority of the bacteria were classified as species ofArthrobacter, Bacillus, Flavobacterium, andXanthomonas. Isolates ofAlcaligenes (Achromobacter), Aeromonas, Chromobacterium, Micrococcus, andPseudomonas were less common. No measurable rates of nitrogen fixation attributable to free-living bacteria were detected by acetylene reduction. Eleven species in six genera of lichens containing a blue-green algal phycobiont showed positive acetylene reduction. One species,Lobaria oregana, accounted for 51% of the total lichen biomass of the canopy. Cellulase and xylanase activity was routinely detected in moss and litter samples, and less frequently in lichens. There was a strong correlation between the two activities for moss (r=0.94) and litter (r=0.81).     

Bacterial and Heterotrophic Nanoflagellate Densities and C‐biomass Estimates Along an Alaskan Tundra Transect with Prediction of Respiratory CO2 Efflux

Although tundra terrestrial ecology is significantly affected by global warming, we know relatively little about how eukaryotic microbial communities respond and how much microbial respiratory CO₂ may be released due to available organic nutrient sources in the permafrost melt. Prior research has shown a strong positive correlation between bacteria and fungi in some Arctic locales; this research focused on the relationships of terrestrial bacteria and heterotrophic nanoflagellates. The densities and estimated C‐biomass of bacteria and heterotrophic nanoflagellates (a major occurring group of protozoa) were assessed in 14 samples obtained along a 10 km transect in northwest AK during the summer of 2012. Two samples were taken, one at the top and one near the base of seven hummocks along the transect. Densities (no./g soil) of bacteria varied from 2.7–16 × 10⁹, and nanoflagellates 0.7–7.9 × 10⁷. C‐biomass (μg/g soil) of bacteria varied from 358 to 2,114, and nanoflagellates 12–37. Additionally, the rate of respiration was analyzed in the laboratory for each soil sample. A linear relationship between soil respiration and bacterial densities was obtained (20 °C): R_s = 12.32 + 14.07 B_d (p ? 0.01); where R_s is soil respiration (nmol/min/g soil) and B_d = bacterial density (no. × 10⁹/g soil).     

The role of water lichens in the biogeochemical processes in the Lake Baikal stony littoral

Water crustose lichens of g. Verrucaria (V. scabra Vězda, V. rheitrophyla Zsch., V. maura Wahlenb. in Ach., and V. hydrela Ach.) and water foliose lichen Collema ramenskii Elenk. were studied in the stony littoral of the western coast of Lake Baikal in 2002–2006. Their densities were highest at depths of 1.5 to 2 m; 95–100% of rock fragments collected from the depths of 1.5–2.2 m were covered by plentiful crustose thalli of Verrucaria spp. Lichens make a large contribute to the destruction of the stony ground in the shallow-water zone. Unlike water, foliose and crustose lichens concentrate Ti, La, Ce, Y, Mn, and Th. Thalli of Collema ramenskii differ from Verrucaria species in their ability to accumulate Zn, Co, and Ni. Compared to the bottom sediments, all the studied water lichen species concentrate Zn, while Collema ramenskiiconcentrate Zn and Mo.     

Environmental factors and ground disturbance affecting the composition of species and functional traits of ground layer lichens on grey dunes and dune heaths of Estonia

A unique, species‐rich and endangered lichen biota can be found on European coastal and inland sand dunes. However, it is increasingly affected by natural succession as well as by anthropogenic disturbances. We studied lichen diversity on the grey dunes and dune heaths of coastal and inland regions of Estonia. A total of 28 study plots were investigated; in each 0.1 ha study plot general environmental variables and anthropogenic disturbances were described and all epigeic lichen species were identified. We found 66 lichenized fungus (lichen) species, including several rare and ten red‐listed lichens. Multivariate analysis (DCA, CCA) was performed to examine gradients in species composition and to relate variation in species data to environmental factors. In addition, we used redundancy analysis (RDA) to relate variation in species’ trait composition to environmental factors. Species composition on grey dunes differed significantly from that on dune heaths. The characteristic species for grey dunes are, besides several Cladonia species, foliose lichens, e.g. Hypogymnia physodes, Parmelia sulcata and Peltigera spp. Also species’ traits composition was different for either habitat, indicating that sorediate lichens, foliose lichens, lichens with cyanobacterium as the main photobiont, and sparsely branched Cladonia species dominate on grey dunes, while esorediate, green‐algal, crustose and richly branched fruticose lichens are common on dune heaths. Soil pH was the most essential environmental variable for determining both species composition and species’ traits composition. The composition of lichen species was also significantly influenced by forest closeness, soil Mg content and cover of bare sand; the effect of ground disturbances was low compared to the effect of these environmental factors. To protect and conserve the species‐rich lichen biota, it is necessary to protect the dune habitats from building activity, to avoid overtrampling in recreation areas and to regularly remove shrubs and trees.     

The vertical gradient of bark pH of twigs and macrolichens in a Picea abies canopy not affected by acid rain

Abstract:The pH of 192 thin, even-aged twigs from 4 height levels of 12 randomly selected trees within a boreal Picea abies canopy naturally exposed to rainfall with a high pH (>5·2) was measured. The largest variation in bark pH was due to the height above the ground. However, a highly significant horizontal variation between trees was also found, apparently due to small-scale soil variations. The biomass of alectorioid lichens increased with increasing height above the ground to at least 12 m, a height interval with fairly constant pH values. The uppermost twigs had an unusually high pH and an abnormal species composition for P. abies, with dominance of the foliose Melanelia exasperatula. The canopy hosted several cyanobacterial lichens, but these were scattered and had low biomass, restricted to lower branches of the trees with the highest bark pH.     

Phagotrophic protists are a key component of microbial communities processing leaf litter under contrasting oxic conditions

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Naked Amoebae in Biofilms Collected from a Temperate Freshwater Pond

Biofilms collected on Plexiglass substrates, from a freshwater pond in northern New York State, were examined microscopically for naked amoebae densities, sizes, diversity, and estimated C‐biomass. Five samples were obtained during summer 2006 and 2007. The densities ranged from 109 to 136/cm² biofilm surface and 285 to 550/mg biofilm dry weight. Sizes ranged from 13 to 200 μm. Diversities ranged from 4.23 to 4.55. C‐biomass ranged from 64 to 543 ng C/cm² and 125 to 1,700 μg C/g dry weight. Thirty morphospecies were identified among the five samples, including very large amoebae in the range of 100–200 μm. Large amoebae (≥ 50 μm) accounted for the largest proportion of the C‐biomass.     

Energetics of Acidianus ambivalens growth in response to oxygen availability

All life requires energy to drive metabolic reactions such as growth and cell maintenance; therefore, fluctuations in energy availability can alter microbial activity. There is a gap in our knowledge concerning how energy availability affects the growth of extreme chemolithoautotrophs. Toward this end, we investigated the growth of thermoacidophile Acidianus ambivalens during sulfur oxidation under aerobic to microaerophilic conditions. Calorimetry was used to measure enthalpy (ΔH_inc) of microbial activity, and chemical changes in growth media were measured to calculate Gibbs energy change (ΔG_inc) during incubation. In all experiments, Gibbs energy was primarily dissipated through the release of heat, which suggests enthalpy‐driven growth. In microaerophilic conditions, growth was significantly more efficient in terms of biomass yield (defined as C‐mol biomass per mole sulfur consumed) and resulted in lower ΔG_inc and ΔH_inc. ΔG_inc in oxygen‐limited (OL) and oxygen‐ and CO₂‐limited (OCL) microaerophilic growth conditions resulted in averages of ?1.44 × 10³ kJ/C‐mol and ?7.56 × 10² kJ/C‐mol, respectively, and average ΔH_inc values of ?1.11 × 10⁵ kJ/C‐mol and ?4.43 × 10⁴ kJ/C‐mol, respectively. High‐oxygen experiments resulted in lower biomass yield values, an increase in ΔG_inc to ?1.71 × 10⁴ kJ/C‐mol, and more exothermic ΔH_inc values of ?4.71 × 10⁵ kJ/C‐mol. The observed inefficiency in high‐oxygen conditions may suggest larger maintenance energy demands due to oxidative stresses and a preference for growth in microaerophilic environments.     

Epiphytic lichen is associated with species richness of gall-inducing aphids but not with niche differentiation among them

The presence of epiphytic foliose lichen amplifies the heterogeneity of habitat by creating shelters for insects living on tree bark. It thus should enhance species number and spatial niche segregation among these canopy insects. We studied this hypothesis in a field experiment using four aphid species that induce galls on Pistacia atlantica trees covered with Xanthoria parietina lichen. In autumn 2008, 3 months after aphid fundatrices were oviposited, we marked six branches on each of 29 trees. Two served as a control, whereas the other four were isolated with insect glue; two of them were scraped with sandpaper to remove epiphytic foliose lichens. We therefore obtained three treatments comprising control branches, isolated branches with lichen, and isolated branches without lichen. In summer 2009, we counted all the galls developing on five new annual shoots on each of 174 branches. We observed more cecidogenic aphid species on all the branches with lichens than without, but each species at different proportions. The different frequencies of utilization of the lichen did not lead to habitat partitioning between species. In conclusion, although habitat heterogeneity itself was associated with species richness and population abundance, it did not induce spatial niche segregation. Considering that many economically important insect species, pests and natural enemies, oviposit or spend some portion of their lives in bark cracks, it is possible that some can use lichens too for protection or/and oviposition sites. As a consequence, lichens may affect management of agrosystems and their impacts should be investigated more deeply in such contexts.     

Diverse Legionella‐Like Bacteria Associated with Testate Amoebae of the Genus Arcella (Arcellinida: Amoebozoa)

Diverse species of Legionella and Legionella‐like amoebal pathogens (LLAPs) have been identified as intracellular bacteria in many amoeboid protists. There are, however, other amoeboid groups such as testate amoeba for which we know little about their potential to host such bacteria. In this study, we assessed the occurrence and diversity of Legionella spp. in cultures and environmental isolates of freshwater arcellinid testate amoebae species, Arcella hemispherica, Arcella intermedia, and Arcella vulgaris, via 16S rRNA gene sequence analyses and fluorescent in situ hybridization (FISH). Analysis of the 16S rRNA gene sequences indicated that A. hemispherica, A. intermedia, and A. vulgaris host Legionella‐like bacteria with 94–98% identity to other Legionella spp. based on NCBI BLAST search. Phylogenetic analysis placed Legionella‐like Arcella‐associated bacteria (LLAB) in three different clusters within a tree containing all other members of Legionella and LLAPs. The intracellular localization of the Legionella within Arcella hosts was confirmed using FISH with a Legionella‐specific probe. This study demonstrates that the host range of Legionella and Legionella‐like bacteria in the Amoebozoa extends beyond members of “naked” amoebae species, with members of the testate amoebae potentially serving an ecological role in the dispersal, protection, and replication of Legionella spp. in natural environments.     

The fate of 15N added to high Arctic tundra to mimic increased inputs of atmospheric nitrogen released from a melting snowpack

Increases in the long‐range aerial transport of reactive N species from low to high latitudes will lead to increased accumulation in the Arctic snowpack, followed by release during the early summer thaw. We followed the release of simulated snowpack N, and its subsequent fate over three growing seasons, on two contrasting high Arctic tundra types on Spitsbergen (79°N). Applications of ¹⁵N (99 atom%) at 0.1 and 0.5 g N m^?2 were made immediately after snowmelt in 2001 as either Na¹⁵NO₃ or ¹⁵NH₄Cl. These applications are approximately 1 × and 5 × the yearly atmospheric deposition rates. The vegetation at the principal experimental site was dominated by bryophytes and Salix polaris while at the second site, vegetation included bryophytes, graminoids and lichens. Audits of the applied ¹⁵N were undertaken, over two or three growing seasons, by determining the amounts of labeled N in the soil (0–3 and 3–10 cm), soil microbial biomass and different vegetation fractions. Initial partitioning of the ¹⁵N at the first sampling time showed that ～60% of the applied ¹⁵N was recovered in soil, litter and plants, regardless of N form or application rate, indicating that rapid immobilization into organic forms had occurred at both sites. Substantial incorporation of the ¹⁵N was found in the microbial biomass in the humus layer and in the bryophyte and lichen fractions. After initial partitioning there appeared to be little change in the total ¹⁵N recovered over the following two or three seasons in each of the sampled fractions, indicating highly conservative N retention. The most obvious transfer of ¹⁵N, following assimilation, was from the microbial biomass into stable forms of humus, with an apparent half‐life of just over 1 year. At the principal site the microbial biomass and vascular plants were found to immobilize the greatest proportion of ¹⁵N compared with their total N concentration. In the more diverse tundra of the second site, lichen species and graminoids competed effectively for ¹⁵NH₄‐N and ¹⁵NO₃‐N, respectively. Results suggest that Arctic tundra habitats have a considerable capacity to immobilize additional inorganic N released from the snow pack. However, with 40% of the applied ¹⁵N apparently lost there is potential for N enrichment in the surrounding fjordal systems during the spring thaw.     

Winter grazing of reindeer in woodland lichen pasture: Effect of lichen availability on the condition of reindeer

Winter grazing of semi-domesticated reindeer (Rangifer t. tarandus) was investigated at the woodland lichen pasture (lichen approximately 550 kg DM ha⁻¹) in Kaamanen, northern Finland during the winter 1996–1997. Nine female reindeer mainly dug their food in the snow for 122 days (3 December–4 April) in a fenced area of 36.3 ha. Over half of the fenced area was lichen dominated dry pine forest. The amount of lichens in lichen forest inside the fence was estimated before and after grazing. Area of grazed and condition of reindeer as well as snow conditions were monitored. Reindeer grazed over the whole area of lichen forest in early winter but from mid-winter they tended to graze on the areas with the greatest lichen abundancy. The amount of lichens measured decreased in the latter areas by 40% and in the other part of the lichen pasture by 17%, respectively. In both of these areas the residual amounts of lichens left after grazing were similar. Of the dominant lichens, the amount of C. stellaris decreased the most and the amount of Cl. uncialis the least. During the study, the estimated average daily area grazed varied from 4 to 87 m² per reindeer. It was calculated that individual reindeer obtained 2.6 kg of lichen DM per day during the most intensive digging period when the body condition score and weight of reindeer increased. Otherwise, the body condition score and weight decreased. Reindeer finished foraging for ground lichens and started to search for arboreal lichens in mid-March when the snow layer was 70–80 cm thick and contained some hard snow layers which lifted reindeer. Both the amount of lichens in the pasture and the snow conditions essentially affect the nutritional status of reindeer in the woodland region during winter. Assuming that a reindeer is able to graze around 30 m² per day in the snow during mid and late winter, there should be, on the basis of energy demand and grazing behaviour of reindeer as well as the nutritive value of lichen, an estimated 1000 kg lichen DM ha⁻¹ available in a good condition woodland lichen pasture.     

Fungal specific primers for PCR‐amplification of mitochondrial LSU in lichens

Fungal specific primer sequences for the amplification of the large subunit of the mitochondrial ribosomal DNA (mtLSU) are presented in this paper. Fungal specific primers make the separation of fungal and algal cells prior to DNA‐extraction from lichens unnecessary. This is especially useful in crustose and small foliose and fruticose lichens. An example from a complex of closely related species of the crustose lichen genus Biatora shows the usefulness of mtLSU‐sequences for studies of infraspecific variability and lower level systematics of lichenized ascomycetes.