Epiphyte Biomass and Canopy Microclimate in the Tropical Lowland Cloud Forest of French Guiana

Recent work on bryophyte diversity in lowland forests of northern South America has suggested the existence of a new type of cloud forest, the ‘tropical lowland cloud forest’ (LCF). LCF occurs in river valleys in hilly areas with high air humidity and morning fog, and is rich in epiphytes. We explored epiphyte abundance and canopy microclimate of LCF in a lowland area (200–400 m asl) near Saül, central French Guiana. We analyzed the vertical distribution of epiphytic cover and biomass on 48 trees, in LCF and in lowland rain forest (LRF) without fog. Trees in LCF had significantly more epiphytic biomass than in LRF; mean total epiphytic biomass in LCF was about 59 g/m², and 35 g/m² in LRF. In all height zones on the trees, total epiphyte cover in LCF exceeded that in LRF, with ca 70 percent mean cover in LCF and ca 15 percent in LRF. During both wet and dry seasons, mean diurnal relative air humidity (RH) was higher in LCF than in LRF, and persistence of high RH after sunrise significantly longer in LCF. We suggest that the prolonged availability of high air humidity in LCF and the additional input of liquid water through fog, enhance epiphyte growth in LCF by shortening the desiccation period and lengthening the period of photosynthetic activity of the plants.     

Influence of management on the species composition of epiphytic cryptogams in Danish Fagus forests

Abstract. The purpose of this study is to evaluate and to quantify the influence of management on the epiphytic species composition in Danish Fagus sylvatica forests. Trunks were chosen at random within selected blocks in both managed and unmanaged Fagus stands.117 trunks in 43 sample plots in three forests were investigated. Cover was estimated for 72 lichens, 29 bryophytes and two fungi on vertical trunks 0.3 ‐2 m above ground. 18 explanatory variables were recorded and arranged in three groups: eight management‐related variables, six micro‐climatic‐ and soil variables, and four geographical‐ and geological variables. Canonical Correspondence Analysis was used to test and to quantify the explaining power of the three groups of variables applying the variation partitioning approach. Detrended Correspondence Analysis was used to evaluate whether important gradients were ignored. It was concluded that the traditional ‘shelterwood uniform system’ used in Fagus forests has a marked influence on the epiphytic species composition. Thus, management‐related variables explained more than one third of the total variation explained, which equals 10.9% of the total variation. Microclimatic and soil variables explained 9.0%, and geographical and geological variables explained 6.0%. The following three recommendations are suggested to reduce the significant influence of management on the epiphytic species composition. First, it should be attempted to ensure a continuous occurrence of some big trees per ha to be left for natural death and decomposition. Second, regular thinning is necessary, especially in the first part of rotation. Third, a multi‐layered forest, where wind and desiccation effects are minimized, is recommended.     

Desiccation Tolerance and Global Change: Implications for Tropical Bryophytes in Lowland Forests

Global change puts an increasing pressure on tropical forests and their inherent diversity by the risk of longer droughts and drier microclimatic conditions within the forest. How organisms will respond is uncertain, especially for organisms highly depending on their microclimatic environment such as bryophytes. An adequate tolerance to desiccation is important to face these changes, however, little is known for tropical bryophytes. We investigated for the first time the desiccation tolerance of epiphytic bryophytes from contrasting microsites at the tropical lowland forest in French Guiana. Using chlorophyll‐fluorescence (F_v/F_m) as an indicator of recovery, we tested: (1) desiccation tolerance for short (3 d) and long (9 d) desiccation events; (2) different desiccation intensities; and (3) recovery by rehydration with water vapor. Species from the canopy were well adapted to desiccation events. Thirteen of 18 species maintained more than 75 percent of their photosynthetic capacity after recovery at the strongest desiccation treatment of 9 d at 43 percent relative humidity (RH). In contrast, species from the understory were sensitive and withstood desiccation only at humid conditions of 75 percent RH and higher. The photosystem of the studied bryophytes was reactivated efficiently in equilibration with water vapor only—a yet neglected phenomenon in bryology. A novel introduced desiccation tolerance index allows global comparison of desiccation tolerances and highlights the sensitivity of understory species. Our results suggest that decreasing humidity caused by climate change and forest degradation could be a concerning threat for understory species.     

Retention of Inorganic Nitrogen by Epiphytic Bryophytes in a Tropical Montane Forest1

We developed and evaluated a model of the canopy of a tropical montane forest at Monteverde, Costa Rica, to estimate inorganic nitrogen (N) retention by epiphytes from atmospheric deposition. We first estimated net retention of inorganic N by samples of epiphytic bryophytes, epiphyte assemblages, vascular epiphyte foliage, and host tree foliage that we exposed to cloud water and precipitation solutions. Results were then scaled up to the ecosystem level using a multilayered model of the canopy derived from measurements of forest structure and epiphyte mass. The model was driven with hourly meteorological and event‐based atmospheric deposition data, and model predictions were evaluated against measurements of throughfall collected at the site. Model predictions were similar to field measurements for both event‐based and annual hydrologic and inorganic N fluxes in throughfall. Simulation of individual events indicated that epiphytic bryophytes and epiphyte assemblages retained 33–67 percent of the inorganic N deposited in cloud water and precipitation. On an annual basis, the model predicted that epiphytic components retained 3.4 kg N ha/yr, equivalent to 50 percent of the inorganic N in atmospheric deposition (6.8 kg N ha/yr). Our results indicate that epiphytic bryophytes play a major role in N retention and cycling in this canopy by transforming highly mobile inorganic N (ca. 50% of atmospheric deposition is NO^?₃) to less mobile (exchangeable NH⁺₄) and recalcitrant forms in biomass and remaining litter and humus.     

Functional diversity of epiphytes in two tropical lowland rainforests, French Guiana: using bryophyte life-forms to detect areas of high biodiversity

Recent studies have described a new tropical lowland forest type in the Guianas, the tropical lowland cloud forest. It is characterized by an enriched epiphytic species diversity particularly for bryophytes compared to common lowland rainforest, and is facilitated by frequent early morning fog events in valley locations. While the increase in epiphytic species diversity in lowland cloud forests has been documented, uncertainties remain as to (1) how this small scale variation in water supply is shaping the functional diversity of epiphytic components in lowland forests, and (2) whether information on functional group composition of epiphytes might aid in discerning these cloud forests from the common lowland rainforest. We compare the distribution of functional groups of epiphytes across height zones in lowland cloud forest and lowland rain forest of French Guiana in terms of biomass, cover as well as the composition of bryophyte life-forms. Both forests differed in functional composition of epiphytes in the canopy, in particular in the mid and outer canopy, with the cloud forest having a higher biomass and cover of bryophytes and vascular epiphytes as well as a richer bryophyte life-form composition. Bryophyte life-forms characteristic for cloud forests such as tail, weft and pendants were almost lacking in the canopies of common rain forest whereas they were frequent in lowland cloud forests. We suggest that ground-based evaluation of bryophyte life-form composition is a straightforward approach for identifying lowland cloud forest areas for conservation, which represent biodiversity hotspots in tropical lowland forests.     

New national and regional bryophyte records, 46

Abstract

Bryophyte biomass and diversity vary strongly with altitude in the tropics. Low abundance and low species numbers in lowland rain forests are most likely due to reduced diurnal activity times combined with high nocturnal respiration rates at high temperatures. This may exclude many montane species from the warm lowlands. However, an alternative hypothesis explains the observed pattern, namely a limited desiccation tolerance of montane species, precipitation being more concentrated but less frequent in most lowland forests compared to montane cloud forests. To test this hypothesis, we studied the desiccation tolerance of four montane and four lowland bryophyte species. The effects of prolonged drought were quantified with chlorophyll fluorescence (F_v/F_m) and the extent of electrolyte leakage. Both montane and lowland species survived dry periods of ≧80 days, which far exceeds the duration of dry periods in the wet lowland tropics. We can thus exclude intolerance to long dry spells as an explaination for the absence of the tested montane species in the lowlands. We should continue to focus on other mechanisms to explain the altitudinal gradient of bryophyte abundance and diversity in the tropics, in order to understand this pattern, as well as to predict future trends under climatic warming.     

Epiphytic ferns and bryophytes of Tasmanian tree‐ferns: A comparison of diversity and composition between two host species

Abstract Ferns, bryophytes and lichens are the most diverse groups of plants in wet forests in south‐eastern Australia. However, management of this diversity is limited by a lack of ecological knowledge of these groups and the difficulty in identifying species for non‐experts. These problems may be alleviated by the identification and characterization of suitable proxies for this diversity. Epiphytic substrates are potential proxies. To evaluate the significance of some epiphytic substrates, fern and bryophyte assemblages on a common tree‐fern species, Dicksonia antarctica (soft tree‐fern), were compared with those on a rare species, Cyathea cunninghamii (slender tree‐fern), in eastern Tasmania, Australia. A total of 97 fern and bryophyte species were recorded on D. antarctica from 120 trunks at 10 sites, and 64 species on C. cunninghamii from 39 trunks at four of these sites. The trunks of C. cunninghamii generally supported fewer species than D. antarctica, but two mosses (particularly Hymenodon pilifer) and one liverwort showed significant associations with this host. Several other bryophytes and epiphytic ferns showed an affinity for the trunks of D. antarctica. Species assemblages differed significantly between both sites and hosts, and the differences between hosts varied significantly among sites. The exceptionally high epiphytic diversity associated with D. antarctica suggests that it plays an important ecological role in Tasmanian forests. Evidently C. cunninghamii also supports a diverse suite of epiphytes, including at least one specialist species.     

哀牢山湿性常绿阔叶林树干4种附生苔藓植物的水分变化对光合、荧光参数的影响

以云南哀牢山湿性常绿阔叶林中的附生苔藓植物为对象,研究林内4种树干附生苔藓植物阿萨羽苔(Plagiochila assamica Steph.)、西南树平藓(Homaliodendron montagneanum(Müll.Hal.)M.Fleisch.)、刀叶树平藓(H.scalpellifolium(Mitt.)M.Fleisch.)、大羽藓(Thuidium cymbifolium(Dozy et Molk.)Dozy et Molk.)的吸水力、失水特征和脱水耐性,稳定碳同位素(δ~(13)C)值,以及水分变化对苔藓净光合速率和荧光参数的影响。结果显示:(1)4种附生苔藓植物均具有较强的吸水能力和较低持水力,脱水耐性较强,脱水后均可迅速复水;(2)4种附生苔藓植物的δ~(13)C值较低,其水分利用效率小于其他维管束植物;除大羽藓外,其余3种附生苔藓雨季的δ~(13)C值与水分利用效率均高于干季;(3)4种附生苔藓植物的净光合速率(Pn)随着含水量的降低而降低,在含水量小于60%~80%时,其最大光化学效率(Fv/Fm)急剧下降,反映出这些附生苔藓植物的光合、荧光特性对水分变化非常敏感。     

Spatial and temporal variation in butterfly biodiversity in a West African forest: lessons for establishing efficient rapid monitoring programmes

Because tropical forests face serious threats and are usually situated in developing countries, cheap and easy Rapid Biodiversity Assessment (RBA) seems an ideal strategy for documenting changes in habitat quality and setting conservation priorities. Fruit‐feeding butterflies are often used as focal taxa to document biodiversity trends in tropical forests. However, the choice of inappropriate sampling strategies may nullify its value. More extensive data sets that sample multiple months and seasons and incorporate the vertical dimension are needed as a baseline for designing reliable sampling regimes. Therefore, we performed a butterfly trapping study in Ghana, using both understorey and canopy traps, and sampling seven periods covering dry and wet seasons. Both individual numbers and species richness were on average three to four times higher in the understorey than in the canopy traps with strikingly different species composition (only 11% overlap in species). The number of species and individual abundance recorded in a month varied extensively. This study underlined the importance of taking into account temporal variation and vertical stratification when designing RBAs of fruit‐feeding butterflies in West African forests. We recommend the use of both understorey and canopy traps and inclusion of both ‘wet’ and ‘dry’ seasons into RBA sampling regimes.     

辽东山地古石河冰缘地貌森林生态系统苔藓物种多样性研究

基于9个20 m×30 m森林群落样地的调查数据,采用物种丰富度、α和β多样性指数,对辽东山地古石河冰缘地貌不同林型石生、树生苔藓植物物种多样性进行定量研究,采用皮尔逊相关分析方法对其影响因素进行分析。结果显示,古石河冰缘地貌苔藓植物共有26科46属59种;不同林型石生、树生苔藓植物物种丰富度和α多样性指数均为:暗针叶林针阔混交林落叶阔叶林;石生苔藓植物β多样性指数最高为落叶阔叶林-针阔混交林间(0.44),最低为落叶阔叶林-暗针叶林间(0.33);树生苔藓植物β多样性指数最高为针阔混交林-暗针叶林间(0.40),最低为落叶阔叶林-暗针叶林间(0.25);分析表明,林冠层郁闭度、海拔高度是影响辽东山地古石河冰缘地貌森林生态系统苔藓物种多样性的重要因子。     

The British Species of the Plagiothecium denticulatum–P. silvaticum Group

Abstract

Anomodon viticulosus was subjected to several different patterns of alternating wet and dry periods, and assimilation was measured at intervals over 49 days. Short periods of desiccation intercalated between 6-day moist periods had little effect on subsequent performance. Short moist periods breaking otherwise continuous desiccation had a very pronounced effect in mitigating desiccation damage. The results provide some evidence for a fairly slight, but probably cumulative, specific deleterious effect of drying and remoistening.     

Old trees contribute bio-available nitrogen through canopy bryophytes

Symbiotic cyanobacteria??bryophyte associations on the forest floor are shown to contribute significantly to stand-level nitrogen budgets through the process of biological nitrogen fixation (BNF), but few studies have considered the role of canopy bryophytes. Given the high biomass of epiphytic bryophytes in many tree species of the North American temperate rain forest, we suggest that canopy bryophytes may contribute substantially to stand-level N dynamics. We confirm the presence of cyanobacteria and measure rates of BNF at three heights (0, 15 and 30 m) in Sitka spruce trees across three watershed estuaries of Clayoquot Sound, British Columbia, Canada. This study is the first to report BNF by cyanobacteria associated with epiphytic and forest floor bryophytes in the coastal temperate rain forest of North America. Cyanobacteria density was significantly greater in epiphytic bryophytes compared to mosses on the forest floor, and rates of BNF were highest at 30 m in the canopy. The majority of total stand-level BNF (0.76 kg N · ha^-1 · yr^-1) occurs in the canopy, rather than on the forest floor (0.26 kg N · ha^-1 · yr^-1). We suggest that BNF by cyanobacterial-bryophyte associations in the canopy of coastal temperate rain forests is a unique source of ecosystem N, which is dependent on large, old trees with high epiphytic bryophyte biomass.     

Mussaenda epiphytica sp. nov. (Rubiaceae), an epiphytic shrub from cloud forest of the Bakossi Mts,western Cameroon

Mussaenda epiphytica Cheek is described as a new species, remarkable for being a cloud forest canopy epiphytic shrub in a genus of terrestial climbers and straggling shrubs. Attention is drawn to its adaptations to the epiphytic habit. The species appears to be endemic to the submontane evergreen forests of the Bakossi Mts, including Mt Kupe, in Cameroon. Its conservation status is assessed as ‘Vulnerable’ using IUCN criteria.     

Response of epiphytic bryophytes to simulated N deposition in a subtropical montane cloud forest in southwestern China

A field manipulation experiment was conducted in a subtropical montane cloud forest in southwestern China to determine the possible responses of epiphytic bryophytes to increasing nitrogen (N) deposition from community to physiology level, and to find sensitive epiphytic bryophytes that may be used as indicators for assessing the degree of N pollution. N addition had significantly negative effects on species richness and cover of the epiphytic bryophyte community. Harmful effects of high N loads were recorded for chlorophyll, growth, and vitality of the species tested. The decline of some epiphytic bryophytes may result from detrimental effects on degradation to photosynthetic pigments. Bazzania himalayana (Mitt.) Schiffn., Bazzania ovistipula (Steph.) Mizut., and Homaliodendron flabellatum (Sm.) Fleisch. are candidates in atmospheric nitrogen monitoring. Epiphytic bryophytes in the montane cloud forest are very sensitive to increasing N deposition and often difficult to recover once they have been destroyed, providing early detection of enhanced N pollution for trees or even the whole forest ecosystem. The inference that increasing N pollution may lead to loss of biodiversity is a concern to the developing economy in western China, and should alert the government to the adverse impacts caused by increased industrial pollution during the process of China’s West Development.     

Benchmarks and predictors of coarse woody debris in native forests of eastern Australia

Fallen coarse woody debris (CWD) is critical to forest biodiversity and function. Few studies model factors that influence CWD availability, although such investigations are critically needed to inform sustainable forest management. We assess benchmark levels of CWD in unharvested native forests and those harvested for timber, across a range of forests in north‐east New South Wales, Australia. We found timber‐harvesting was the dominant driver of CWD, with almost double the count (pieces ha^?1) and volume (m³ ha^?1) of total CWD in selectively harvested than unharvested sites. This pattern was consistent across wet and dry forest types. Harvested sites had greater counts of hollow‐bearing logs, and greater volumes of small and medium‐sized CWD (15–50 cm diameter) than unharvested sites. There was no effect of harvesting on the volume of large CWD (>51 cm diameter). Total volumes of CWD (>15 cm diameter) varied from 114 to 166 m³ ha^?1. We found few differences in CWD counts and volumes between forest types, with grassy woodlands and forests containing less CWD than other dry and shrubby forest types, reflecting lower potential input rates. The CWD levels recorded here are similar to those recorded in dry and wet sclerophyll forests elsewhere in Australia and are typical of global estimates for ‘old growth’ forests. Using general linear models we captured up to 57% of the variation in CWD across sites, and found that timber harvesting, topography and the numbers of standing hollow‐bearing and dead trees were significant predictors of CWD. Values for unharvested forest provide a benchmark that could be used to inform retention guidelines for CWD in managed forests in this region. Further assessment of the effect of repeat timber harvesting is needed to fully understand its impact on CWD dynamics, especially if forest residues resulting from timber harvesting are removed from native forests for bioenergy production.     

865. NEPENTHES DUBIA

The history, taxonomic placement, relationships, mode of trapping and cultivation of Nepenthes dubia Danser, an epiphytic fly‐eater from the cloud forests of the Bukit Barisan range of the west coast of Sumatra, are discussed. This is one of the eight species of Sect. Montanae Danser with ‘egg‐cup’ like pitchers. A key is provided to the eight species and they are formally named as Nepenthes Sect. Montanae subsect. Poculae‐ovis.     

Environmental flow allocations in monsoonal Hong Kong

1. Hong Kong streams are subject to aggressive water extractions but the downstream water needs of ecosystems – i.e. environmental flow (e‐flow) requirements – have not yet been addressed. This study investigated hydro‐ecological relationships that could be used to establish e‐flow allocations for streams in monsoonal Hong Kong. 2. Data were collected during the wet and dry seasons from 10 unpolluted streams experiencing a gradient of flow reductions (c. 0–98%). Relationships between flow conditions (percentage discharge reduction and absolute discharge volume) and responses of macroinvertebrate composition and periphyton condition were established for each season. 3. Declines in richness of Ephemeroptera and abundance of hydropsychid caddisflies, as well as increases in the proportion of predators, were linearly related to percentage discharge reduction during both seasons. Relationships were also recorded for eight other macroinvertebrate richness or compositional metrics during the dry season only. Relationships between macroinvertebrate assemblage attributes and absolute discharge volume across downstream reaches were also evident. Periphyton was relatively insensitive to flow reductions and did not provide useful hydro‐ecological relationships, although declines in autotrophic index were related to percentage discharge reduction during the dry season. 4. Using hydro‐ecological relationships established for macroinvertebrates, two levels of e‐flow were proposed: a ‘threshold’ intended to maintain near‐natural conditions and a ‘degradation limit’ that allowed no more than 25% of the maximum indicator response to flow reduction. Calculated threshold e‐flows required downstream allocation of ≥74% of natural flows; degradation limit e‐flows were ≥12% (wet) and ≥27% (dry). The discharge needed to maintain threshold conditions was 30–105 L s^?1 (wet) and 5–14 L s^?1 (dry), with degradation limit e‐flows of 19–57 L s^?1 (wet) and 3–6 L s^?1 (dry), relative to natural mean discharges of 77–303 L s^?1 (wet) and 3–18 L s^?1 (dry). 5. The proposed e‐flow allocations are indicative only, and significant obstacles to implementation have yet to be surmounted. Any such implementation requires monitoring of outcomes in order to refine the allocations and inform adaptive flow management for Hong Kong streams.     

Bryophyte phytomass in tropical ecosystems

Knowledge about bryophyte production and phytomass in the tropics is poor and has been derived principally from studies of epiphytic bryophytes. Such work has been undertaken in Transvaal, Tanzania, Venezuela, Peru and Borneo. Except for the studies in Peru and Borneo, the vegetation types studied and the methods used have been too variable to allow many comparisons or generalizations to be made. However, in general it can be stated that phytomass of epiphytic bryophytes in equatorial latitudes increases from the tropical lowlands to the foreslline. This striking phenomenon has been hypothetically explained by gradients of factors such as precipitation, humidity, temperature and desiccation, or combinations of these. Recent experiments on the gas exchange of tropical-montane bryophytes have revealed that the rate of net assimilation decreases dramatically above 25^oC and that high temperature combined with low light intensities, as realized in the tropical lowland forest, results in high rates of respiration and insufficient net photosynthesis. Experiments with temperate bryophytes show the same results. If can be assumed that tropical lowland species have special physiological adaptations to survive these unfavourable conditions, such as a specialized phytochrome system or effective storage of photosynthetic products. Additional factors could include the relatively high nutrient supply provided by abundant precipitation in tropical montane forests, and damage to cells by strong desiccation in the tropical lowlands.     

Effects of intermittent desiccation on bryophytes

Abstract

(1) Gas exchange measurements on Rhytidiadelphus loreus and Anomodon viticulosus following re-moistening after a period of desiccation showed an initial stimulation of respiration, followed immediately by progressive recovery of photosynthesis within the first 1–6 h. A. viticulosus after 48 h dry reached the compensation point within a few minutes of re-moistening; R. lore us after 7 days dry reached the compensation point after about 4 hours.

(2) Hylocomium splendens, R. loreus and Tortula ruraliformis were subjected to the following patterns of alternating wet and dry periods over a period of 18 weeks: continuous dry, 1 day wet/6 days dry, 6 days wetll day dry, 1 day wet/1 day dry, 7 days wet/7 days dry, continuous wet.

(3) Only T. ruraliformis still attained a positive net assimilation rate after the continuous dry treatment for the duration of the experiment. All three species retained a positive net assimilation rate throughout after all the remaining treatments.

(4) A moist period of 24 h allowed substantially complete recovery from the after-effects of preceding desiccation.

(5) Experiments in which two dry periods were separated by moist periods of 6 h, 24 hand 7 days showed that recovery in H. splendens and R. loreus was incomplete in 6 h but confirmed that it was complete in 24 h.

(6) H. splendens showed constant or slightly increased desiccation-tolerance in successive dry periods. R. loreus showed a well marked increase in tolerance (hardening) following a previous dry/wet cycle.

(7) The effect of intermittent desiccation can be predicted approximately from the successive effect of the individual dry periods.

(8) It is doubtful whether there is any specific deleterious effect of drying and re-moistening; any such effect that does exist can only be small.     

Resurrection kinetics of photosynthesis in desiccation‐tolerant terrestrial green algae (Chlorophyta) on tree bark

The rough bark of orchard trees (Malus) around Darmstadt is predominantly covered in red to purple‐brown layers (biofilms) of epiphytic terrestrial alga of Trentepohlia umbrina. The smooth bark of forest trees (Fagus sylvatica L. and Acer sp.) in the same area is covered by bright green biofilms composed of the green algae Desmococcus, Apatococcus and Trebouxia, with a few cells of Coccomyxa and ‘Chlorella’ trebouxioides between them. These algae are desiccation tolerant. After samples of bark with the biofilms were kept in dry air in darkness for various periods of time, potential quantum yield of PSII, F_v/F_m, recovered during rehydration upon rewetting. The kinetics and degree of recovery depended on the length of time that the algae were kept in dry air in the desiccated state. Recovery was better for green biofilm samples, i.e. quite good even after 80 days of desiccation (F_v/F_m = ca. 50% of initial value), than the red samples, where recovery was only adequate up to ca. 30–40 days of desiccation (F_v/F_m = ca. 20–55% of initial value). It is concluded that the different bark types constitute different ecophysiological niches that can be occupied by the algae and that can be distinguished by their capacity to recover from desiccation after different times in the dry state.