The weathering action of saxicolous lichens in maritime Antarctica

Summary Xanthoria elegans (Link) Th Fr. and Lecidea lapicida (Ach.) were studied on volcanic andesite, and Rhizocarpon geographicum (L.) DC. and Bacidia stipata Lamb on a volcanigenic sediment, using light microscopy, infrared spectroscopy, X-ray diffraction and transmission electron microscopy. Feldspars were present in the rocklichen interface to a lesser extent than in the underlying rock. R. geographicum was found to alter the minerals in the rock on which it grew without producing any new minerals in the rock/lichen interface, in contrast to the observations for this species on granite in temperate regions. Beneath of the thallus of L. lapicida there was calcium oxalate and some micas of the illite type, which may have been degradation products of various phyllosilicates in the rock. B. stipata, an endemic Antarctic lichen, had the greatest capacity to weather the rock and had weddellite (dihydrate calcium oxalate) and calcite in the contact area as well as many bacteria. The presence of crystalline oxalate, imogolite, allophane, carbonates (calcite) and amorphous material not found in the parent rock indicates biomineralization processes attributable to the lichens.     

Growth and vitality of epiphytic lichens

 We tested the hypothesis that changed microclimate at induced forest edges causes reduced growth of epiphytic lichens. Two foliose, green algal lichens were transplanted to the lower canopy of a mature Picea abies forest at six distances (2, 6.25, 12.5, 25, 50 and 100 m) from a clearcut. The biomass growth in Platismatia glauca (6.2% in 16 months) was 41% higher than in Lobaria pulmonaria (4.4%). We found no growth reduction near the forest edge. In contrast, the highest growth in both species occurred within 12 m from the edge. Further, fluorescence and chlorophyll measurements showed that lichen vitality was unaffected by distance from edge. The light intensity was 4.3 times higher at the edge than in the interior during the growing season, but there were only minor differences in air temperature and relative humidity. Monitoring of thallus water content revealed clear differences in both number and length of wetting and drying cycles. However, the total time with water content sufficient for photosynthetic activity was only slightly higher at the edge. The data thus indicate that our gradient in microclimate was too small to significantly affect lichen growth, and that lichens are largely metabolically inactive when large edge-interior contrasts in microclimate occur. Lichen response to forest edge microclimate results from intricate interactions among several biotic and abiotic factors. Linking data on lichen growth, microclimate and thallus water content with physiological measurements provides a framework for future studies of the mechanisms behind abiotic edge effects. Received: 15 April 1996 / Accepted: 21 June 1996     

亚热带山地森林附生地衣的移植生长及其对环境变化的响应

本研究选取云南哀牢山亚热带森林系统6种常见的附生大型地衣进行为期2年的移植实验,分析其在原生林林外、林缘以及林内3种生境下的生物量增长速率和健康率等的差异,并探讨其生长对环境因子变化的响应。结果显示,网肺衣Lobaria retigera和平滑牛皮叶Sticta nylanderiana在林缘处生物量增长最快;而槽枝Sulcaria sulcata、多花松萝Usnea florida、皮革肾岛衣Nephromopsis pallescens、针芽肺衣Lobaria isidiophora 4种地衣则都是在林外光照更强、湿度更低的条件下生长得最好,林内最差。除网肺衣外,所有地衣的健康率均在林外较好,林缘次之,林内最差。非参数相关分析表明,6种地衣的生长速率和光照以及温度呈正相关,与大气湿度呈负相关;温度和光照是影响地衣生长最主要的生境因子。     

Iron and phosphate uptake explains the calcifuge–calcicole behavior of the terricolous lichens Cladonia furcata subsp. furcata and C. rangiformis

Mechanisms causing the calcifuge–calcicole behavior of lichens are largely unexplored. Studying the case examples of two closely related terricolous lichens, the calcifuge Cladonia furcata subsp. furcata and the calcicole C. rangiformis, we found that preference for acidic or calcareous soils in these lichens is related to iron and phosphate uptake as in vascular plants. In laboratory studies, the calcicole species was more efficient in the intracellular uptake of Fe³⁺ and phosphate at pH 8 than the calcifuge species. At pH 3, intracellular uptake of Fe²⁺ in the calcicole species significantly exceeded that in the calcifuge species suggesting that calcicole lichens suffer from toxicity symptoms by excess Fe²⁺ at acidic sites. Though these observations parallel findings from calcifuge and calcicole vascular plants, mechanisms leading to the different iron and phosphate uptake characteristics in the studied calcifuge and calcicole lichens may differ from those in vascular plants and should be the topic of future research. A role of the depside atranorin in facilitating iron uptake by reducing Fe³⁺ in the apoplast is hypothesized.     

Cellular injuries in epiphytic lichens transplanted to air polluted areas

The epiphytic lichens Hypogymnia physodes (L.) W. Wats, and Bryoria capillaris (Ach.) Brodo & D. Hawksw. growing on spruce branches were transplanted from a clean rural area to the environment of a fertilizer plant and a pulp mill in central Finland. The common major pollutants in these environments are SO₂, NO_x and ammonia but the fertilizer plant also emits fluorides. In the transmission electron microscope two main types of cellular injuries were observed both in algal and fungal cells in both species and in both environments. The first type, characterized by rapid degeneration of cell organelles, was an apparently acute injury leading to plasmolysis and rapid death of the cells. The second type was considered chronic injury and involved changes in chloroplast shape, swelling of mitochondria and increased density of cytoplasm in algal cells, and increased vacuolization and appearance of dark vacuolar accumulations in fungal cells. The cytoplasmic storage droplets decreased gradually in size both in algae and fungi. The acute injury was mainly seen in the lichens transplanted to sites with higher pollution levels near the sources, and was more usual in algal than in fungal cells. B. capillaris was more susceptible to acute injury than H. physodes. An additional injury type was detected in algal cells of both lichen species in the vicinity of the fertilizer plant. This type was characterized by severe swelling of thylakoids and their interspaces and granulation of thylakoid membranes, and was suspected to be related to the effects of fluorides. The injuries as seen in light microscope, and expressed as increased proportion of dead algae and visible bleaching of thallus, were usually observable simultaneously. The injuries as seen in the electron microscope always preceded other injuries; they were clearly observable already after one week of transplantation in the more polluted sites but developed more slowly in the less polluted ones. The time lack between these injuries and the visible ones was 2–3 weeks in the more polluted sites, but several months or even years in the less polluted ones.     

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。     

Introduction of saxicolous lichens distributed in coastal rocks of U-do islet in Jeju, Korea

This study reports, for the first time, the investigation of the distribution of Korean saxicolous lichens in the coastal rocks of U-do islet, which is known as an unpolluted zone in Jeju. More than thirty lichens were obtained and investigated from the coastal rocks frequently contacted by seawater. A molecular analysis using PCR amplification of the rRNA ITS regions revealed the coastal rock lichens could be placed into 8 families and 14 genera, Ramalinaceae (Bacidia, Ramalina), Physciaceae (Buellia, Dirinaria, Phaeophyscia, Physcia, Pyxine), Lecanoraceae (Candelaria, Lecanora), Parmeliaceae (Xanthoparmelia), Graphidaceae (Graphis), Pertusariaceae (Pertusaria), Rhizocarpaceae (Rhizocarpon), and Teloschistaceae (Caloplaca), showing a diversity of lichens, with foliose (flat leaf-like), crustose (crust-like), and fruticose (miniature shrub-like) life forms might be distributed in the coastal rocks. These findings suggested the possibility that the lichens identified in the present work might be resistant to a salty environment.     

亚热带森林附生地衣压力-体积曲线分析及其适用性

附生地衣是森林生态系统中的重要组成部分, 在维护森林物种多样性以及水分和养分循环等方面发挥着重要作用。地衣类群的水势特征以及压力-体积(PV)曲线及相关参数是否适用于探索地衣等变水性生物类群应对水分胁迫机制, 仍有待阐明。该研究以中国西南地区云南哀牢山亚热带山地森林中5个类群15种常见附生地衣为对象, 探讨了其PV曲线及水势参数的功能群和物种水平的变化。结果显示, 蓝藻型地衣的内部含水量(WC_internal)和共质体水含量(Rs)显著高于绿藻型地衣, 是其1倍以上, 但其他参数表现出较高的一致性。功能群水平上, 地衣仅WC_internal、Rs和膨压损失点的相对含水量(RWC_TLP)具有显著性差异; 但物种间差异均非常显著。结合主成分分析进一步发现, PV曲线及相关参数在评估地衣整体应对水分胁迫以及揭示生境水分条件选择策略时具有较大的限制性, 受到共生藻和生长型的显著影响; 但内部最大持水力可以解释蓝藻型地衣的生境适应策略, 而部分参数如饱和渗透势(Ψsat)和RWC_TLP可以分别用于解释狭叶地衣、阔叶地衣和枝状地衣对生境水分条件的适应。研究表明PV曲线及水势参数不适用于地衣类群整体的抗旱性评价, 和其他生物类群进行抗旱性比较时更要慎用。     

Organic and inorganic nitrogen uptake in lichens

In order to learn more about nitrogen (N) acquisition in lichens, and to see whether different lichens differ in their affinity to various N sources, N uptake was measured in 14 various lichen associations (species). These species represented various morphologies (fruticose or foliose), contrasting microhabitat preferences (epiphytic or terricolous), and had green algal, cyanobacterial or both forms of photobionts. N was supplied under non-limiting conditions as an amino acid mixture, ammonium, or nitrate, using ¹⁵N to quantify uptake. Carbonyl cyanide m-chlorophenylhydrazone (CCCP) was used to separate active and passive uptake. Thallus N, amino acids, soluble polyol concentrations, and the biont-specific markers chlorophyll a and ergosterol were quantified, aiming to test if these metabolites or markers were correlated with N uptake capacity. Ammonium uptake was significantly greater and to a higher extent passive, relative to the other two N sources. Nitrate uptake differed among lichen photobiont groups, cyanobacterial lichens having a lower uptake rate. All lichens had the capacity to assimilate amino acids, in many species at rates equal to nitrate uptake or even higher, suggesting that organic N compounds could potentially have an important role in the N nutrition of these organisms. There were no clear correlations between N uptake rates and any of the measured metabolites or markers. The relative uptake rates of ammonium, nitrate and amino acids were not related to morphology or microhabitat.Abbreviations CCCP Carbonyl cyanide m-chlorophenylhydrazone - Chl Chlorophyll - N Nitrogen     

Stand characteristics in colour-infrared aerial photographs as indicators of epiphytic lichens

Information about forest biodiversity has so far been collected mostly by using field inventories, but it is desirable to find other methods that can cover large areas at a lower cost. In a forest landscape covering 2000 ha in southern Sweden we tested if colour-infrared (CIR) aerial photographs on the scale of 1:30000 can be used to interpret forest stand characteristics correlated to the occurrence of epiphytic lichens that are Red-listed or otherwise indicate high nature conservation value (signal species). Using logistic regression we found that the interpreted stand variables tree height and crown structure class were significantly correlated to the occurrence of Red-listed species. For signal species, the variables tree height, percentage of southern deciduous trees and crown structure class were significantly correlated to the occurrence. The logistic regression models successfully predicted a significantly higher probability to find Red-listed species in the stands that actually contained such species compared to stands without Red-listed species. The same was true for stands with signal species. We conclude that interpretation of CIR aerial photographs could be a useful method to find certain groups of epiphytic lichens in surveys covering large areas.     

Water relations and carbon dioxide exchange of epiphytic lichens in the Namib fog desert

Although there is only negligible rainfall, frequent nocturnal fog, dew and high air humidity support a luxurious lichen vegetation in the coastal zone of the central Namib Desert (Namibia). In earlier publications, we have studied ecophysiological performance of a series of epilithic and terrestrial lichens. Here, we have extended this work to three epiphytic species (Heterodermia namaquana, Ramalina lacera, and Xanthoria turbinata) that inhabit the sparse perennial shrubs growing in this area. Our intention, monitoring lichen CO₂ exchange, their water relations and microclimate conditions, was to determine the functional mechanisms that allow these epiphytes to exist under the special conditions of a fog desert. Measurements were conducted mainly during the spring season.The epiphytic lichens showed response patterns very similar to the epilithic and epigaeic species at the same site. Their metabolism was activated through moistening by dew and/or fog during the night and, in the very early morning, they exhibited the typical brief peak of net photosynthesis (NP) between sunrise and desiccation. The thalli were almost completely dry for the remainder of the day. Average duration of the positive NP during the morning peak was about 3 h. Dew condensation, alone, resulted in activation that provided 58–63% of integrated carbon income (ΣNP) as compared to fog (plus dew). In the late afternoon, there was a tendency for hydration to increase again, due to water vapour uptake at higher air humidity, and this allowed on some days a brief additional period of very low rates of photosynthesis shortly before sunset.Light response of photosynthesis showed “sun-plant” characteristics with saturation around 1000 μmol m⁻² s⁻¹ photosynthetically active photon flux density (PPFD). Light compensation point (LCP) of CO₂ exchange after sunrise was highly dependent on actual water content (WC) for X. turbinata: at low hydration it was ca. 10 μmol m⁻² s⁻¹ PPFD whilst, at high WC, it was almost 80 μmol m⁻² s⁻¹ PPFD. In contrast, LCP of R. lacera was almost independent of WC. This phenomenon was probably due to differences in thallus structure.Maximal attained NP and daily ΣNP both showed a saturation-type response to previous maximal nocturnal WC. Neither parameter was increased substantially when higher maximal thallus WCs were produced by experimental moistening in the night. All three species, despite their different morphologies, performed optimally at the highest nocturnal moistening achieved by natural fog and were not able to make use of higher hydration.The three studied epiphytes were similar in their chlorophyll-related rates of NP. Due to lower chlorophyll content, dry weight and carbon-related NP of X. turbinata was only about one-third of that of the other two species. The average carbon income on days with fog and/or dew hydration during the spring season amounted to 2.4 and 2.1 mgC (gC)⁻¹ day⁻¹ (related to thallus carbon content) for H. namaquana and R. lacera, respectively. This primary production was of similar magnitude to those found for the terrestrial species at the same site.     

High cyanobiont selectivity of epiphytic lichens in old growth boreal forest of Finland

Here, cyanobiont selectivity of epiphytic lichen species was examined in an old growth forest area in Finland. Samples of the eight lichen species were collected from the same aspen (Populus tremula) and adjacent aspens in the same stand. The cyanobionts of these samples were compared with free and symbiotic Nostoc obtained from other habitats and geographic regions. Our results, based on the phylogenetic analysis of a partial small subunit of the ribosomal DNA (16S rDNA) and the rbcLX gene complex did not show any correlation with the geographic origin of the samples at any spatial scale. Instead, there was a correlation between the cyanobionts and the alleged taxonomy of their mycobionts. The results indicate that the lichen species examined are highly selective towards their cyanobiont partners. Only Lobaria pulmonaria proved to be more flexible, being able to associate with a wide range of Nostoc. A same Nostoc strain was found to form associations with taxonomically unrelated lichens indicating that the cyanobiont-mycobiont associations as a whole were not highly specific in the examined species.     

The influence of environmental factors on the spatial distribution of saxicolous lichens in a Norwegian coastal community

Abstract. The effects of vegetation cover, radiation, micro‐habitat variables and maritime influence on the floristic composition of a saxicolous community in Vingen, western Norway were studied. Particular emphasis is put on the local distribution of Fuscidea cyathoides, Ochrolechia tartarea, Ophioparma ventosa and Pertusaria corallina. Very little of the variation in the lichen community composition is directly related to measured micro‐environmental variables but variance partitioning shows that vegetation cover explains more of the floristic variation than radiation, maritime influence and microhabitat variables. Logistic regression analyses nevertheless indicate that the micro‐environment influences the spatial distribution of the four species. The high fraction of unexplained floristic variation, 91%, is suggested to result from (1) lack of fit of data to the response model; (2) some influential environmental variables that have not been recorded; (3) local historical factors that affect present day distribution and/or (4) apparent randomness in colonization. The results also agree with the view that the four lichen species in this study are able to co‐exist in the long‐term because of different spatial distributions resulting from different strategies with respect to ecology, dispersion and interaction.     

Enhanced phosphate uptake and polyphosphate accumulation in Burkholderia cepacia grown under low pH conditions

Of bacterial cells in a sample of activated sludge, 34% contained detectable intracellular polyphosphate inclusions following Neisser staining, when grown on glucose/mineral salts medium at pH 5.5; at pH 7.5 only 7% of cells visibly accumulated polyphosphate. In a sludge isolate of Burkholderia cepacia chosen for further study, maximal removal of phosphate and accumulation of polyphosphate occurred at pH 5.5; levels were up to 220% and 330% higher, respectively, than in cells grown at pH 7.5. During the early stationary phase of growth at pH 5.5 a maximum level of intracellular polyphosphate that comprised 13.6% of cellular dry weight was reached. Polyphosphate kinase activity was detected in actively growing cells only when cultured at pH 5.5. The phenomenon of acid-stimulated phosphate uptake and polyphosphate accumulation in this environmental bacterial population parallels observations previously made by us in the yeast Candida humicola and may thus represent a widespread microbial response to low external pH values.     

Highly efficient uptake of phosphorus in epiphytic bromeliads

Background and Aims

Vascular epiphytes which can be abundant in tree crowns of tropical forests have to cope with low and highly intermittent water and nutrient supply from rainwater, throughfall and stem flow. Phosphorus rather than nitrogen has been suggested as the most limiting nutrient element, but, unlike nitrogen, this element has received little attention in physiological studies. This motivated the present report, in which phosphate uptake kinetics by leaves and roots, the subsequent distribution within plants and the metabolic fate of phosphate were studied as a step towards an improved understanding of physiological adaptations to the conditions of tree canopies.

Methods

Radioactively labelled [³²P]phosphate was used to study uptake kinetics and plant distribution of phosphorus absorbed from bromeliad tanks. The metabolism of low molecular phosphorus metabolites was analysed by thin-layer chromatography followed by autoradiography.

Key Results

Uptake of phosphate from tanks is an ATP-dependent process. The kinetics of phosphorus uptake suggest that epiphytes possess effective phosphate transporters. The K_m value of 1·05 µm determined for leaves of the bromeliad Aechmea fasciata is comparable with values obtained for the high affinity phosphate transporters in roots of terrestrial plants. In this species, young leaves are the main sink for phosphate absorbed from tank water. Within these leaves, phosphate is then allocated from the basal uptake zone into distal sections of the leaves. More than 80 % of the phosphate incorporated into leaves is not used in metabolism but stored as phytin.

Conclusions

Tank epiphytes are adapted to low and intermittent nutrient supply by different mechanisms. They possess an effective mechanism to take up phosphate, minimizing dilution and loss of phosphorus captured in the tank. Available phosphorus is taken up from the tank solution almost quantitatively, and the surplus not needed for current metabolism is accumulated in reserves, i.e. plants show luxury consumption. Young, developing leaves are preferentially supplied with this nutrient element. Taken together, these features allow epiphytes the efficient use of scarce and variable nutrient supplies.Key words: Epiphytic bromeliads, phosphorus uptake, forest canopies, luxury consumption, phytotelms, plant nutrition, Aechmea fasciata     

Distribution pattern of air pollution and epiphytic lichens in the Osaka Plain (Japan)

The geographic distributions of two common species of epiphytic lichens,Phaeophyscia limbata (Poelt) Kashiw. andLecanora pulverulenta Müll. Arg., on city park trees in the Osaka Plain and their relation to the distribution of SO₂ and NO₂ was examined. The concentrations of SO₂ and NO₂ are greater in the central area of this plain, and decrease with distance from the center. The frequency ofL. pulverulenta is lowest in the central area, and increases gradually toward the periphery. In contrast, the frequency ofP. limbata is highest in the intermediate area, and lower in the central and peripheral areas, i.e. at intermediate SO₂ and NO₂ concentrations. We conclude thatL. pulverulenta is more useful thanP. limbata as a bioindicator of air pollutants such as SO₂ and NO₂.     

Vertical distribution of epiphytic bryophytes and lichens emphasizes the importance of old beeches in conservation

During storms in 2005, a number of beech trees fell over at Biskopstorp, SW Sweden, offering the opportunity to study epiphytes along entire stems. In total 16 beech trees in four beech stands representing three different age classes were included. For each tree, 2 m segments from the base to the top were surveyed. In total 115 species were found (76 lichens, 39 bryophytes), of which 30 were considered to be of conservation concern (22 lichens, 8 bryophytes). For lichens significantly more species were recorded above 2 m in height, whereas more bryophytes were recorded below 2 m in height. Certain red-listed lichens were recorded only above 2 m in height on old trees. In a second data set from the same area 140 age-determined beech trees were surveyed for species of conservation concern at the heights 0–2 and 2–5 m, respectively. These species were found almost exclusively on old beech trees, and presence at 2–5 m was recorded, with one exception, only on those trees which also had species of conservation concern at 0–2 m. Records of these species correlated significantly to microhabitat variables, i.e. the presence of rough bark and moss cover higher up the stems on the old trees. This study indicates that surveying only the base in really old beech forests can underestimate both the number of species of conservation concern and their population sizes. However, surveys restricted to the base in rather even-aged beech stands catch a large proportion of the trees with species of conservation concern.