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。     

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。     

不同水分条件下三种附生地衣的光合作用特性

附生地衣是哀牢山湿性常绿阔叶林生态系统中重要的结构性组分。通过对该区域山地森林中3种典型附生地衣平滑牛皮叶 (Sticta nylanderiana)、网肺衣 (Lobaria retigera) 和橄榄斑叶 (Cetrelia olivetorum)在不同水分条件下的光合光响应及荧光参数的测定分析,结果显示,附生地衣光补偿点 (LCP)、光饱和点 (LSP)较高,对强光适应能力较强。在水分胁迫 (含水量5%~10%) 条件下,3种附生地衣的最大净光合速率 (Pmax) 仅为17~50nmol·g-1·s-1。随着含水量的增加,地衣的最大净光合速率 (Pmax) 与暗呼吸速率 (Rday) 逐渐增大,LCP降低,而LSP随之提高,这表明3种附生地衣具备“阳生植物”的某些特性,从而能够在一定程度上适应野外光照较强的灌丛、向阳林冠等生境。地衣叶绿素光反应中心初始荧光参数 (F0) 和最大光化学效率 (Fv/Fm) 随含水量下降而显著降低,暗示其光反应中心对水分有很强的敏感性。水分条件的改善有助于附生地衣的光反应中心进入到较高的生理活性状态。     

Bole epiphytic lichens as potential indicators of environmental change in subtropical forest ecosystems in southwest China

Lichen epiphytes are applied as excellent environmental indicators worldwide. However, very little is known about epiphytic lichen communities and their response to forest dynamics in subtropical China. This paper proposes the applications of the cover, diversity, and functional traits of epiphytic lichens to assess environmental changes associated with succession in subtropical forests of southwest China. Bole lichens were sampled from 120 plots of eight representative forest types in the Ailao Mountains. Total cover, species richness, diversity and community structure of bole lichens differed significantly among forest types, and the highest cover and diversity occurred in the Populus bonatii secondary forest (PBSF). Sixty-one indicator species were associated with particular forest types and more than 50% occurred in the PBSF. Both cover and diversity of most lichen functional groups varied regularly during forest succession. Lichen pioneer species were not displaced by competitively superior species as succession proceeds and cyanolichens were more prevalent in secondary forests. The results also highlight the importance of habitat variables such as canopy openness, host diversity, forest age, tree size, the size of the largest tree, tree density, and basal area on the lichen community. Consequently, our findings support the notion that epiphytic lichens, in terms of cover, diversity, species composition and functional traits can be used as effective indicators for large-scale and long-term forest monitoring. More importantly, the narrowly lobed foliose group was the best candidate indicator of environmental conditions in this region. The combined application of lichen indicator species and functional groups seemed to be a more reliable and more powerful method for monitoring forest dynamics in subtropical montane ecosystems.     

Predicting lichen hydration using biophysical models

Two models for predicting the hydration status of lichens were developed as a first step towards a mechanistic lichen productivity model. A biophysical model included the water potential of the air, derived from measurements of air temperature, relative humidity and species-specific rate constants for desiccation and rehydration. A reduced physical model, included only environmental parameters, assuming instantaneous equilibration between the lichen and the air. These models were developed using field and laboratory data for three green algal lichens: the foliose epiphytic Platismatia glauca (L.) W. Culb., the fruticose epiphytic Alectoria sarmentosa (Ach.) Ach. and the fruticose, terricolous and mat-forming Cladina rangiferina (L.) Weber ex Wigg. The models were compared and validated for the same three species using data from a habitat with a different microclimate. Both models predicted the length and timing of lichen hydration periods, with those for A. sarmentosa and P. glauca being highly accurate—nearly 100% of the total wet time was predicted by both the biophysical and physical models. These models also predicted an accurate timing of the total realized wet time for A. sarmentosa and P. glauca when the lichens were wet. The model accuracy was lower for C. rangiferina compared to the epiphytes, both for the total realized wet time and for the accuracy of the timing for the hydration period. These results demonstrate that the stochastic and continually varying hydration status of lichens can be simulated from biophysical data. Further development of these models to also include water-related activity, light and temperature conditions during the hydration events will then be a potent tool to assess potential lichen productivity in landscapes and habitats of various microclimatic conditions.     

亚热带常绿阔叶林附生地衣凋落物的物种多样性和生物量的边缘效应

地衣是亚热带山地森林系统附生生物类群的重要组成部分之一,对环境变化极其敏感。为了更好地了解附生地衣对森林边缘效应的响应,我们在云南哀牢山地区原生山地常绿阔叶林中,分别在距林缘5m、25m、50m和 100m处设立样地,收集附生大型地衣的凋落物1年;分析附生地衣凋落物的物种多样性和生物量、功能群特征和组成结构对林缘深度变化的响应特征。研究结果显示,边缘效应能够显著提高林缘附生地衣群落的物种多样性和生物量;其发生的距离最深不超过25m。林缘-林内梯度上,不同地衣功能群的响应模式具有各自的特征。排序分析表明仅在5m样地与其他样地之间存在显著差异,指示种分析则发现仅5m样地具有指示种。哀牢山原生林中边缘效应促进林缘附生地衣生长和分布的现象,可能与当地高湿环境削减了地衣的高光损伤以及以叶状和灌状类群为主的地衣个体受到风力破坏的程度相对较低有关。     

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.     

Modeling study on photosynthetic-light response curves of a C4 plant,maize

Aims A light response curve can reflect a plant’s ability to utilize light, which is also a key tool in determining the relationship between photosynthetic capacity and environmental factors; however the model accuracies concerning the light response curve remain elusive. The objectives of this study were to compare and assess the model accuracies related to a light response curve and the effects of drought. Methods A field rain shelter was used to control the soil water conditions. To obtain photosynthesis parameters from the light response curve and the drought effects, the relevant models (including the rectangular model, non-rectangular hyperbolic model, modified rectangular hyperbolic model, exponential model, quadratic function model, and a newly modified model) were applied to fit the light response curves. The validity of each model was tested by analyzing the differences between the fitted values obtained by the models and the measured values. Important findings The newly modified model has been proved to performing relatively better in accurately describing the light response curve patterns, and credibly obtaining the crucial photosynthetic parameters such as the maximum net photosynthetic rate, light saturation point, light compensation point, and dark respiration rate, especially under high radiation conditions.     

Associations between corticolous microarthropod communities and epiphytic cover on bark

The discontinuous bark cover formed by epiphytic lichens and algae provides a mosaic of microhabitats for the fauna. Multivariate analyses applied to 1800 samples collected in Belgian Lorraine (southern Belgium) during each of the four seasons has made it possible to distinguish five major classes of arthropod microcommunities. Two of them are confined to special habitats or places at certain seasons, viz.

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  a Pseudochermes fraxini (Homoptera) community found on Fraxinus during the summer, and "trophically" different from others;
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  a Vertagopus arborea (Collembola) community observed in foliose lichens in St. Mard mainly in winter.

The three other classes are directly related to the epiphytic cover, viz.

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  a Dometorina plantivaga (Oribatida) community found in crustose epiphytes;
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  an Eueremaeus oblongus/Trichoribates trimaculatus (Oribatida) community sheltered by foliose lichens;
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  an Entomobrya nivalis (Collembola)/Cerobasis guestfalicus (Psocoptera) community observed in fruticose lichens.

The ecological meaning of those microcommunities (mosaic and stratification patterns, seasonal variation, succession) is discussed. The results support the hypothesis that corticolous microcoenoses are associated with the epiphyte type and that their composition is greatly affected by the vegetation stratification pattern on bark.     

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.     

Non-invasive monitoring of photosynthetic activity and water content in forest lichens by spectral reflectance data and RGB colors from photographs

There is a need for non-invasive monitoring of temporal and spatial variation in hydration and photosynthetic activity of red-listed poikilohydric autotrophs. Here, we simultaneously recorded kinetics in RGB-colors (photos), reflectance spectra, water content, maximal (F_V/F_M), and effective quantum yield of PSII (Φ_PSII) during desiccation in foliose lichens differing in cortical characteristics and photobionts. The spectral absorbance peaks of chlorophyll a, phycocyanin, and phycoerythrin were clearly displayed at high hydration levels. Brightness and total RGB colors of the lichens strongly increased during desiccation. The normalized difference vegetation index (NDVI) efficiently estimated hydration level and Φ_PSII – a proxy for lichen photosynthesis – in all species, including threatened old forest lichens. Color and reflectance indices based on green wavelengths gave good estimates of water content in cephalo- and chlorolichens, but not in cyanolichens with a wider range of photosynthetic pigments. Due to species-specific characteristics, species-wise calibration is essential for non-invasive assessments of lichen functioning.     

Will forest conservation areas protect functionally important diversity of fungi and lichens over time?

Incorporating functional values in biodiversity monitoring systems could add novel perspectives of the status of biodiversity in conservation areas. Stable frequencies of large foliose nitrogen-fixing cyanolichens likely have positive effects on the nitrogen budget of forests and provide food, material and shelter for invertebrates, gastropods and birds. Stable volumes of deadwood and frequencies of associated fungi provide an important supporting function for ecosystem services such as nutrient cycling, carbon storage and soil formation. Based on regional monitoring data from boreal old-growth forest nature reserves and key habitats, we tested for changes in the frequency of various functionally important substrates and species over time. We detected significant reductions in the frequency of indicator cyanolichens occurring on deciduous trees already after 10 years in key habitats, despite non-significant changes in their host substrates. Frequencies of indicator pendulous lichens Alectoria sarmentosa and Bryoria nadvornikiana had also decreased in key habitats, despite overall stable volumes of large conifer host trees. Lichen reductions were more pronounced in the smaller key habitats compared to the larger formally protected nature reserves, likely due to degrading fragmentation and isolation effects. In contrast to these lichens, the average frequencies of old-growth forest indicator fungi decaying coniferous deadwood and common fungi on deciduous trees (Fomes fomentarius) and coniferous trees (Fomitopsis pinicola) remained unchanged. The studied cyanolichens and fruiting fungi generally had similar extinction rates over 10 years, whilst only cyanolichens had substantially lower colonization rates. Amid a severely fragmented landscape, conservation areas seem to struggle in preserving some of the basic old-growth forest values.     

Photosynthesis in chlorolichens: the influence of the habitat light regime

The hypothesis that CO₂ gas exchange and chlorophyll a fluorescence (ChlaF) of lichens vary according to the light regimes of their original habitat, as observed in vascular plants, was tested by analysing the photosynthetic performance of 12 populations of seven dorsoventral, foliose lichens collected from open, south-exposed rocks to densely shaded forests. Light response curves were induced at optimum thallus water content and ChlaF emission curves at the species-specific photon flux at which the quantum yield of CO₂ assimilation is the highest and is saturating the photosynthetic process. Photosynthetic pigments were quantified in crude extracts. The results confirm that the maximum rate of gross photosynthesis is correlated with the chlorophyll content of lichens, which is influenced by light as well as by nitrogen availability. Like leaves, shade tolerant lichens emit more ChlaF than sun-loving ones, whereas the photosynthetic quantum conversion is higher in the latter.     

Effects of groundwater salinity on the characteristics of leaf photosynthesis and stem sap flow in Tamarix chinensis

AimsThe objective of this study was to investigate the change pattern of leaves photosynthesis and stem sap flow of Tamarix chinensisin under different groundwater salinity, which can be served as a theoretical basis and technical reference for cultivation and management of T. chinensis in shallow groundwater table around Yellow River Delta.MethodsThree-year-old T. chinensis, one of the dominated species in Yellow River Delta, was selected. Plants were treated by four different salinity concentrations of groundwater—fresh water (0 g∙L^-1), brackish water (3.0 g∙L^-1), saline water (8.0 g∙L^-1), and salt water (20.0 g∙L^-1) under 1.2 m groundwater level. Light response of photosynthesis and the diurnal courses of leaf transpiration rate, stem sap flux velocity and environment factors under different groundwater salinity were determined via LI-6400XT portable photosynthesis system and a Dynamax packaged stem sap flow gauge based on stem-heat balance method, respectively.Important findings The result showed that groundwater salinity had a significant impact on photosynthesis efficiency and water consumption capacity of T. chinensis by influencing the soil salt. The net photosynthetic rate (P_n), maximum P_n, transpiration rate, stomatal conductance, apparent quantum yield and dark respiration rate increased first and then decreased with increasing groundwater salinity, while the water use efficiency (WUE) continuously decreased. The mean P_n under fresh water, brackish water and salt water decreased by 44.1%, 15.1% and 62.6%, respectively, compared with that under saline water (25.90 µmol∙m^-2∙s^-1). The mean WUE under brackish water, saline water and salt water decreased by 25.0%, 29.2% and 41.7%, respectively, compared with that under fresh water (2.40 µmol∙mmol^-1). With the increase of groundwater salinity from brackish water to salt water, light saturation point of T. chinensisdecreased while the light compensation point increased, which lead to the decrease of light ecological amplitude and light use efficiency. Fresh water and brackish water treatment helped T. chinensis to use low or high level light, which could significantly improve the utilization rate of light energy. The decrease in P_n of T. chinensis was mainly due to non-stomatal limitation under treatment from saline water to fresh water, while the decrease in P_n of T. chinensis was due to stomatal limitation from saline water to salt water. With increasing groundwater salinity, stem sap flux velocity of T. chinensis increased firstly and then decreased, reached the maximum value under saline water. The mean stem sap flux velocity under fresh water, brackish water and salt water decreased by 61.8%, 13.1% and 41.9%, respectively, compared with that under saline water (16.96 g·h^-1). Tamarix chinensis had higher photosynthetic productivity under saline water treatment, and could attained high WUE under severe water deprivation by transpiration, which was suitable for the growth of T. chinensis.     

Photosynthesis of the cyanobacterial soil-crust lichen Collema tenax from arid lands in southern Utah, USA: role of water content on light and temperature responses of CO2 exchange

1. The gelatinous cyanobacterial Collema tenax is a dominant lichen of biotic soil crusts in the western United States. In laboratory experiments, we studied CO₂ exchange of this species as dependent on water content (WC), light and temperature. Results are compared with performance of green-algal lichens of the same site investigated earlier.
2. As compared with published data, photosynthetic capacity of C. tenax is higher than that of other cyanobacterial and green-algal soil-crust species studied. At all temperatures and photon flux densities of ecological relevance, net photosynthesis (NP) shows a strong depression at high degrees of hydration; maximal apparent quantum-use efficiency of CO₂ fixation is also reduced. Water requirements (moisture compensation point, WC for maximal NP) are higher than that of the green-algal lichens. Collema tenax exhibits extreme 'sun plant' features and is adapted to high thallus temperatures.
3. Erratic rain showers are the main source of moisture for soil crusts on the Colorado Plateau, quickly saturating the lichens with liquid water. High water-holding capacity of C. tenax ensures extended phases of favourable hydration at conditions of high light and temperature after the rain for substantial photosynthetic production. Under such conditions the cyanobacterial lichen appears superior over its green-algal competitors, which seem better adapted to habitats with high air humidity, dew or fog as prevailing source of moisture.     

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.     

Effects of drought-rewatering-drought on photosynthesis and growth of maize

Aims Our objective was to investigate the responses of maize photosynthesis and growth to repeated drought.Methods Maize seedlings were exposed to different soil water deficit for three weeks, then rewatering for one week, and again to different water deficit for three weeks, to examine the effects of repeated drought on photosynthesis and growth.Important findings After the first water deficit treatments, under severe drought, plant height, total leaf area of individual plant, shoot and root biomass declined significantly, also transpiration rate (T_r), stomatal conductance (G_s), intercellular CO₂ concentration (C_i), net photosynthetic rate (P_n), maximum net photosynthetic rate (A_max), but light compensation point and dark respiration rate increased significantly. Under medium drought, plant height, leaf area, and shoot biomass decreased significantly, but root biomass did not vary, hence, the ratio of roots to shoots (R/S) increased. Moreover, plants did not show significant differences in photosynthetic parameters. After rewatering, photosynthesis and growth rate of plants previously exposed to water deficit could recover to the levels of well-watered plants, but plant height and leaf area did not recover to the levels of the control. When maize were subjected to recurrent drought, plants pre-exposed to medium drought showed no significant difference in plant height, biomass, and photosynthetic parameters, but a significant decrease in leaf area, compared to plants only exposed to second medium drought. Plants pre-exposed to severe drought had significantly higher T_r, G_s, C_i, P_n, A_max, and, apparent quantum yield but significantly lower plant height, leaf area, and biomass than plants without previous exposure. These results indicated that the first severe drought significantly reduced photosynthetic capacity and maize growth, rewatering could recover photosynthesis and growth rate to the levels of well-watered plants, but could not eliminate the adverse influence of the first drought on growth. The first medium drought could stimulate the growth of maize root system and significantly increased R/S, which can enhance maize drought resistance to subsequent repeated drought, and maintain the total biomass in the control level; the first severe drought could enhance maize drought resistance to subsequent repeated drought in the aspect of photosynthesis, but could not compensate for the adverse effect of early drought on plant growth. Hence, in practice, drought hardening should be limited in the level of medium drought, and avoiding severe drought.     

Phosphorus Fertilization Increases the Abundance and Nitrogenase Activity of the Cyanolichen Pseudocyphellaria crocata in Hawaiian Montane Forests

Nitrogen-fixing epiphytes (especially lichens with a cyanobacterial symbiont—cyanolichens) have the potential to contribute significant amounts of nitrogen (N) to montane tropical forests, which are typically low in N—but the factors controlling the abundance and distribution of epiphytic cyanolichens are poorly understood. In long-term fertilization experiments in montane forests on a 4.l million-yr-old Oxisol on the island of Kauà'i and on a 152-yr-old lava flow on the island of Hawai'i, the epiphytic cyanolichen Pseudocyphellaria crocata increased significantly in abundance in canopies of host trees fertilized with phosphorus (P). There was no response to fertilization with N or other essential elements. Nitrogen-fixation rates were also elevated in lichens in P-fertilized plots at both sites. Phosphorus supply to host trees may be an important factor controlling N inputs to montane tropical forests by N-fixing epiphytes.     

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.