Hemiepiphytic figs kill their host trees: acquiring phosphorus is a driving factor.

Hemiepiphytic figs killing their host trees is an ecological process unique to the tropics. Yet the benefits and adaptive strategies of their special life history remain poorly understood. We compared leaf phosphorus (P) content data of figs and palms worldwide, and functional traits and substrate P content of hemiepiphytic figs (Ficus tinctoria), their host palm and nonhemiepiphytic conspecifics at different growth stages in a common garden. We found that leaf P content of hemiepiphytic figs and their host palms significantly decreased when they were competing for soil resources, but that of hemiepiphytic figs recovered after host death. P availability in the canopy humus and soil decreased significantly with the growth of hemiepiphytic figs. Functional trait trade-offs of hemiepiphytic figs enabled them to adapt to the P shortage while competing with their hosts. From the common garden to a global scale, the P competition caused by high P demand of figs may be a general phenomenon. Our results suggest that P competition is an important factor causing host death, except for mechanically damaging and shading hosts. Killing hosts benefits hemiepiphytic figs by reducing interspecific P competition and better acquiring P resources in the P-deficient tropics, thereby linking the life history strategy of hemiepiphytic figs to the widespread P shortage in tropical soils.

A combination of morphological and photosynthetic functional traits maintains the vertical distribution of bryophytes in a subtropical cloud forest.

PREMISE: The distribution and performance of bryophyte species vary with vertical gradients, as a result of changes in environmental factors, especially light. However, the morphological and physiological drivers of bryophyte distribution along forest vertical gradients are poorly understood. METHODS: For 18 species of mosses and liverworts distributed among three vertical microhabitats (ground, tree trunk, and branch, variance in 28 morphological and photosynthetic functional traits was comparatively analyzed among the microhabitats and bryophyte life-forms in a subtropical cloud forest in Ailao Mountain, Yunnan, southwestern China. Principal component analysis (PCA) was used to summarize trait differences among bryophyte species. RESULTS: In contrast to trunk and ground dwellers, branch dwellers tended to reduce light interception (smaller leaf and cell sizes, lower chlorophyll content), protect against damage from intense irradiation (higher ratios of carotenoids to chlorophyll), raise light energy use (higher photosynthetic capacity), and cope with lower environmental moisture (pendant life-forms, thicker cell walls). The PCA showed that ecological strategies of bryophytes in response to levels of irradiation were specialized in branch dwellers, although those of ground and trunk dwellers were less distinct. CONCLUSIONS: Environmental filtering shaped the combination of functional traits and the spatial distribution of bryophytes along the vertical gradients. Bryophyte species from the upper canopy of cloud forests show narrow variation in functional traits in high-light intensity, whereas species in the lower vertical strata associated with low-light intensity used contrasting, but more diverse ecological strategies.

When facilitation meets clonal integration in forest canopies.

Few studies have explored how - within the same system - clonality and positive plant-plant interactions might interact to regulate plant community composition. Canopy-dwelling epiphytes in species-rich forests provide an ideal system for studying this because many epiphytic vascular plants undertake clonal growth and because vascular epiphytes colonize canopy habitats after the formation of nonvascular epiphyte (i.e. bryophyte and lichen) mats. We investigated how clonal integration of seven dominant vascular epiphytes influenced inter-specific interactions between vascular epiphytes and nonvascular epiphytes in a subtropical montane moist forest in southwest China. Both clonal integration and environmental buffering from nonvascular epiphytes increased survival and growth of vascular epiphytes. The benefits of clonal integration for vascular epiphytes were higher when nonvascular epiphytes were removed. Similarly, facilitation from nonvascular epiphytes played a more important role when clonal integration of vascular epiphytes was eliminated. Overall, clonal integration had greater benefits than inter-specific facilitation. This study provides novel evidence for interactive effects of clonality and facilitation between vascular and nonvascular species, and has implications for our understanding of a wide range of ecosystems where both high levels of clonality and facilitation are expected to occur.

Ecological stoichiometry of the epiphyte community in a subtropical forest canopy.

Epiphytes in tree canopies make a considerable contribution to the species diversity, aboveground biomass, and nutrient pools in forest ecosystems. However, the nutrient status of epiphytes and their possible adaptations to nutrient deficiencies in the forest canopy remain unclear. Therefore, we analyzed the stoichiometry of five macroelements (C, N, P, K, and Ca) in four taxonomic groups (lichens, bryophytes, ferns, and spermatophytes) to investigate this issue in a subtropical montane moist evergreen broad-leaved forest in Southwest China. We found that the interspecific variations in element concentrations and mass ratios were generally greater than the intraspecific variations. And there were significant stoichiometric differences among functional groups. Allometric relationships between N and P across the epiphyte community indicated that P might be in greater demand than N with an increase in nutrients. Although canopy nutrients were deficient, most epiphytes could still maintain high N and P concentrations and low N:P ratios. Moreover, ferns and spermatophytes allocated more limited nutrients to leaves than to stems and roots. To alleviate frequent drought stress in the forest canopy, vascular epiphytes maintained several times higher K concentrations in their leaves than in the tissues of lichens and bryophytes. Our results suggest that epiphytes may have evolved specific nutrient characteristics and adaptations, so that they can distribute in heterogeneous canopy habitats and maintain the stability of nutrient metabolism.

National surveillance of 2,3,7,8-substituted polychlorinated dibenzo-p-dioxins/furans in soil in Taiwan.

In this study, the polychlorinated dibenzo-p-dioxin/furan (PCDD/F) levels in 381 soil samples coming from different background areas (n = 238) and contaminated areas (n = 143) in Taiwan were investigated from 2011 to 2015 using high resolution gas chromatograph/high resolution mass spectrometry (HRGC/HRMS). The contaminated areas showed higher PCDD/F contamination as compared to the background areas (1230 vs 749 pg/g dry weight (dw)); 14.0 vs 6.25 pg WHO2005-TEQ/g dw). The lowest levels were recorded in the outlying islands (1.28 pg WHO2005-TEQ/g dw). PCDD/F congener distribution profiles in the background and contaminated areas as well as on the islands varied. OCDD was the dominant congener in almost all locations, including the northern background (87.7%) and central contaminated (74.2%) areas. Other dominant species included OCDF and 1,2,3,4,6,7,8-HpCDD. Levels of PCDD/F-TEQs in Taiwanese soils, including the background areas, were higher than in some developing countries and regions with global background levels due to high industrialization except for the contaminated areas. Geographic differences in dioxin-contaminated soils were also shown in this study. Higher soil dioxin-TEQs were observed in locations with high populations and population densities. Despite the limitations, the economic status of Taiwan represented by the annual averaged family income (AAFI) was positively correlated to soil dioxin-TEQs.

Epiphytic bryophytes as bio-indicators of atmospheric nitrogen deposition in a subtropical montane cloud forest: Response patterns, mechanism, and critical load.

Increasing trends of atmospheric nitrogen (N) deposition due to pollution and land-use changes are dramatically altering global biogeochemical cycles. Bryophytes, which are extremely vulnerable to N deposition, often play essential roles in these cycles by contributing to large nutrient pools in boreal and montane forest ecosystems. To interpret the sensitivity of epiphytic bryophytes for N deposition and to determine their critical load (CL) in a subtropical montane cloud forest, community-level, physiological and chemical responses of epiphytic bryophytes were tested in a 2-year field experiment of N additions. The results showed a significant decrease in the cover of the bryophyte communities at an N addition level of 7.4 kg ha-1 yr-1, which is consistent with declines in the biomass production, vitality, and net photosynthetic rate responses of two dominant bryophyte species. Given the background N deposition rate of 10.5 kg ha-1yr-1 for the study site, a CL of N deposition is therefore estimated as ca. 18 kg N ha-1 yr-1. A disordered cellular carbon (C) metabolism, including photosynthesis inhibition and ensuing chlorophyll degradation, due to the leakage of magnesium and potassium and corresponding downstream effects, along with direct toxic effects of excessive N additions is suggested as the main mechanism driving the decline of epiphytic bryophytes. Our results confirmed the process of C metabolism and the chemical stability of epiphytic bryophytes are strongly influenced by N addition levels; when coupled to the strong correlations found with the loss of bryophytes, this study provides important and timely evidence on the response mechanisms of bryophytes in an increasingly N-polluted world. In addition, this study underlines a general decline in community heterogeneity and biomass production of epiphytic bryophytes induced by increasing N deposition.

Physiological Responses of Two Epiphytic Bryophytes to Nitrogen, Phosphorus and Sulfur Addition in a Subtropical Montane Cloud Forest.

Atmospheric depositions pose significant threats to biodiversity and ecosystem function. However, the underlying physiological mechanisms are not well understood, and few studies have considered the combined effects and interactions of multiple pollutants. This in situ study explored the physiological responses of two epiphytic bryophytes to combined addition of nitrogen, phosphorus and sulfur. We investigated the electrical conductivity (EC), total chlorophyll concentration (Chl), nutrient stoichiometry and chlorophyll fluorescence signals in a subtropical montane cloud forest in south-west China. The results showed that enhanced fertilizer additions imposed detrimental effects on bryophytes, and the combined enrichment of simulated fertilization exerted limited synergistic effects in their natural environments. On the whole, EC, Chl, the effective quantum yield of photosystem II (ΦPSII) and photochemical quenching (qP) were the more reliable indicators of increased artificial fertilization. However, conclusions on nutrient stoichiometry should be drawn cautiously concerning the saturation uptake and nutrient interactions in bryophytes. Finally, we discuss the limitations of prevailing fertilization experiments and emphasize the importance of long-term data available for future investigations.

Organic nitrogen uptake is a significant contributor to nitrogen economy of subtropical epiphytic bryophytes.

Without any root contact with the soil, epiphytic bryophytes must experience and explore poor, patchy, and heterogeneous habitats; while, the nitrogen (N) uptake and use strategies of these organisms remain uncharacterized, which obscures their roles in the N cycle. To investigate the N sources, N preferences, and responses to enhanced N deposition in epiphytic bryophytes, we carried out an in situ manipulation experiment via the (15)N labelling technique in an Asian cloud forest. Epiphytic bryophytes obtained more N from air deposition than from the bark, but the contribution of N from the bark was non-negligible. Glycine accounted for 28.4% to 44.5% of the total N in bryophyte tissue, which implies that organic N might serve as an important N source. Increased N deposition increased the total N uptake, but did not alter the N preference of the epiphytic bryophytes. This study provides sound evidence that epiphytic bryophytes could take up N from the bark and wet deposition in both organic and inorganic N forms. It is thus important to consider organic N and bark N sources, which were usually neglected, when estimating the role of epiphytic bryophytes in N cycling and the impacts of N deposition on epiphytic bryophytes in cloud forests.

Survival and Growth of Epiphytic Ferns Depend on Resource Sharing.

Locally available resources can be shared within clonal plant systems through physiological integration, thus enhancing their survival and growth. Most epiphytes exhibit clonal growth habit, but few studies have tested effects of physiological integration (resource sharing) on survival and growth of epiphytes and whether such effects vary with species. We conducted two experiments, one on individuals (single ramets) and another on groups (several ramets within a plot), with severed and intact rhizome treatments (without and with physiological integration) on two dominant epiphytic ferns (Polypodiodes subamoena and Lepisorus scolopendrium) in a subtropical montane moist forest in Southwest China. Rhizome severing (preventing integration) significantly reduced ramet survival in the individual experiment and number of surviving ramets in the group experiment, and it also decreased biomass of both species in both experiments. However, the magnitude of such integration effects did not vary significantly between the two species. We conclude that resource sharing may be a general strategy for clonal epiphytes to adapt to forest canopies where resources are limited and heterogeneously distributed in space and time.

Bioaccumulation of mercury and polychlorinated dibenzo-p-dioxins and dibenzofurans in salty water organisms.

Mercury and polychlorinated dibenzo-p-dioxins and dibenzofurans (PCDD/Fs) accumulate in organisms through food webs and exert potentially toxic effects on aquatic organisms and humans. This study examined the levels of mercury and PCDD/Fs in organisms and sediment samples collected from a saltwater pond at the An-Shun site, a chloralkali factory that shut down in Tainan City, Taiwan. It was also a pentachlorophenol production plant. After the factories were shut down in the 1980s, mercury and PCDD/Fs contamination remained, posing severe health hazards. The correlation between PCDD/Fs congener accumulation patterns in distinct fish organs and the sediment was evaluated. Mercury and PCDD/Fs levels in all the fish samples exceeded food safety limits, and the concentrations of mercury and PCDD/Fs in each species were closely correlated (n = 12, Spearman's rank correlation [R] = 0.811, p < 0.01). The mercury concentrations were positively but non-significantly correlated with the weight (n = 11, R = 0.741, p < 0.01) and length (n = 11, R = 0.618, p < 0.05) of the species. The fish likely accumulated the contaminants through ingestion of other organisms or the sediment. However, after the pollutants entered a fish, they exhibited distinct accumulation patterns because of their differing chemical properties. Specifically, the mercury concentration was correlated with organism weight and length, whereas the PCDD/Fs concentration was associated with organ lipid content. The study results are valuable for assessing the health risks associated with ingesting mercury- and PCFF/F-contaminated seafood from the study site.

Higher clonal integration in the facultative epiphytic fern Selliguea griffithiana growing in the forest canopy compared with the forest understorey.

BACKGROUND AND AIMS: The advantage of clonal integration (resource sharing between connected ramets of clonal plants) varies and a higher degree of integration is expected in more stressful and/or more heterogeneous habitats. Clonal facultative epiphytes occur in both forest canopies (epiphytic habitats) and forest understories (terrestrial habitats). Because environmental conditions, especially water and nutrients, are more stressful and heterogeneous in the canopy than in the understorey, this study hypothesizes that clonal integration is more important for facultative epiphytes in epiphytic habitats than in terrestrial habitats. METHODS: In a field experiment, an examination was made of the effects of rhizome connection (connected vs. disconnected, i.e. with vs. without clonal integration) on survival and growth of single ramets, both young and old, of the facultative epiphytic rhizomatous fern Selliguea griffithiana (Polypodiaceae) in both epiphytic and terrestrial habitats. In another field experiment, the effects of rhizome connection on performance of ramets were tested in small (10 × 10 cm(2)) and large (20 × 20 cm(2)) plots in both epiphytic and terrestrial habitats. KEY RESULTS: Rhizome disconnection significantly decreased survival and growth of S. griffithiana in both experiments. The effects of rhizome disconnection on survival of single ramets and on ramet number and growth in plots were greater in epiphytic habitats than in terrestrial habitats. CONCLUSIONS: Clonal integration contributes greatly to performance of facultative epiphytic ferns, and the effects were more important in forest canopies than in forest understories. The results therefore support the hypothesis that natural selection favours genotypes with a higher degree of integration in more stressful and heterogeneous environments.

Water relations and gas exchange of fan bryophytes and their adaptations to microhabitats in an Asian subtropical montane cloud forest.

Fan life forms are bryophytes with shoots rising from vertical substratum that branch repeatedly in the horizontal plane to form flattened photosynthetic surfaces, which are well suited for intercepting water from moving air. However, detailed water relations, gas exchange characteristics of fan bryophytes and their adaptations to particular microhabitats remain poorly understood. In this study, we measured and analyzed microclimatic data, as well as water release curves, pressure-volume relationships and photosynthetic water and light response curves for three common fan bryophytes in an Asian subtropical montane cloud forest (SMCF). Results demonstrate high relative humidity but low light levels and temperatures in the understory, and a strong effect of fog on water availability for bryophytes in the SMCF. The facts that fan bryophytes in dry air lose most of their free water within 1 h, and a strong dependence of net photosynthesis rates on water content, imply that the transition from a hydrated, photosynthetically active state to a dry, inactive state is rapid. In addition, fan bryophytes developed relatively high cell wall elasticity and the osmoregulatory capacity to tolerate desiccation. These fan bryophytes had low light saturation and compensation point of photosynthesis, indicating shade tolerance. It is likely that fan bryophytes can flourish on tree trunks in the SMCF because of substantial annual precipitation, average relative humidity, and frequent and persistent fog, which can provide continual water sources for them to intercept. Nevertheless, the low water retention capacity and strong dependence of net photosynthesis on water content of fan bryophytes indicate a high risk of unbalanced carbon budget if the frequency and severity of drought increase in the future as predicted.

Combination of a fast cleanup procedure and a DR-CALUX® bioassay for dioxin surveillance in Taiwanese soils.

Our goal was to determine dioxin levels in 800 soil samples collected from Taiwan. An in vitro DR-CALUX® assay was carried out with the help of an automated Soxhlet system and fast cleanup column. The mean dioxin level of 800 soil samples was 36.0 pg-bioanalytical equivalents (BEQs)/g dry weight (d.w.). Soil dioxin-BEQs were higher in northern Taiwan (61.8 pg-BEQ/g d.w.) than in central, southern, and eastern Taiwan (22.2, 24.9, and 7.80 pg-BEQ/g d.w., respectively). Analysis of multiple linear regression models identified four major predictors of dioxin-BEQs including soil sampling location (ß = 0.097, p < 0.001), land use (ß = 0.065, p < 0.001), soil brightness (ß = 0.170, p < 0.001), and soil moisture (ß = 0.051, p = 0.020), with adjusted R2 = 0.947 (p < 0.001) (n = 662). An univariate logistic regression analysis with the cut-off point of 33.4 pg-BEQ/g d.w. showed significant odds ratios (ORs) for soil sampling location (OR = 2.43, p < 0.001), land use (OR = 1.47, p < 0.001), and soil brightness (OR = 2.83, p = 0.009). In conclusion, four variables, including soil sampling location, land use, soil brightness, and soil moisture, may be related to soil-dioxin contamination. Soil samples collected in northern Taiwan, and especially in Bade City, soils near industrial areas, and soils with darker color may contain higher dioxin-BEQ levels.

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.

Regeneration pattern of primary forest species across forest-field gradients in the subtropical mountains of Southwestern China.

Evergreen broad-leaved forest is now gradually degraded and fragmented, and there is an increase in the amount of habitat edges as a result of long-term human activity. However, the role of edges in the regeneration of primary forest species is poorly understood. After 20 years of the edge creation, we analyzed primary forest species distribution and abundance, and changes in floristic composition, vegetation structure across forest-field gradients in Ailao Mountain, SW China. Our results revealed that there was a higher abundance and richness of primary species, late secondary species and thorny lianas at the distances 0-50 m than at the distances more than 50 m from the edge into the forest exterior. At the distances >50 m, no individuals of dominant canopy trees Lithocarpus xylocarpus, Castanopsis wattii, and L. jingdongensis were found, whereas the abundance of early pioneer shrub species and herbaceous cover was significantly greater. The richness of primary species showed a decrease with increasing distances from the forest edge to the exterior, particularly of medium-seeded primary species showing a drastic decrease. Moreover, no large-seeded primary species occurred at the distances >60 m. This study indicates that the forest edge as a buffer zone may be in favor of primary species regeneration. A dense shrub and herb layer, and seed dispersal may be the major factors limiting the forest regeneration farther from the forest edge. Therefore, to facilitate forest recovery processes, management should give priority to the protection of buffer zones of this forest edge.

[Microbial community and its activities in canopy- and understory humus of two montane forest types in Ailao Mountains, Northwest China].

Mid-montane moist evergreen broadleaved forest (MMF) and top-montane dwarf mossy forest (DMF) are the two major natural forest types in subtropical mountainous area of Ailao Mountains, Northwest China. In this paper, a comparative study was made on the microbial composition, quantity, biochemical activity, metabolic activity, and their seasonal dynamics in the canopy- and understory humus of the two forest types. The composition, quantity, and metabolic activity of the microbes in the canopy humus of dominant tree species in MMF and DMF were also analyzed. In the canopy humus of the two forest types, the amounts of fungi and actinomycetes, microbial biomass C and N, and intensities of nitrogen fixation and cellulose decomposition were significantly higher than those in understory humus. Meanwhile, the amount of cellulose-decomposing microbes (ACDM), cellulose decomposition intensity, microbial biomass C and N, and metabolic activity in the canopy humus of MMF were significantly higher than those of DMF. The amounts of bacteria, fungi, and aerobic nitrogen-fixing bacteria (ANFB) and the metabolic activity in the canopy humus of MMF and DMF were significantly higher in wet season than in dry season, while a contradictory trend was observed on the amount of actinomycetes. No significant difference was observed on the amount of ACDM between wet season and dry season. For the two forest types, the amounts of microbes and their biochemical activities in canopy humus had a larger seasonal variation range than those in understory humus. There was a significant difference in the amounts of the microbes in canopy humus among the dominant tree species in MMF and DMF, especially in wet season. The microbes in canopy humus played important roles in maintaining the biodiversity of epiphytes in the canopy, and in supplying the needed nutrients for the vigorous growth of the epiphytes.

[Ecological adaptability of different provenance Eupatorium adenophorum Spreng].

By using reciprocal transplant method, Eupatorium adenophorum Spreng. seedlings of five different provenances were reciprocally transplanted into six different sites in Yunnan Province, with the survival rate, plant height, branch number, biomass, flower number per plant, and seed production measured during the period from April 2007 to May 2008. The seedlings growth and reproductive traits of different provenance E. adenophorum all showed strong plasticity to environmental conditions. With increasing latitude and altitude of transplant site, plant height, branch number, biomass, flower number per plant, and seed production of different provenance E. adenophorum decreased, and the differences in these growth and reproductive traits were significant among the transplant sites. However, there were no significant differences in these traits among different provenance E. adenophorum. Provenance and transplant site had no significant interactive effects on the above-mentioned traits except seed production. At each site, the survival rate, growth potential, and reproductive capability of local provenance E. adenophorum didn't have any superiority, illustrating that the success in the invasion of E. adenophorum in Yunnan Province was mainly due to the phenotypic plasticity of the plant, while local adaptability only played lesser important role.

[Composition and carbon storage of woody debris in moist evergreen broad-leaved forest and its secondary forests in Ailao Mountains of Yunnan Provinve].

This paper studied the composition and carbon storage of woody debris in the primary moist evergreen broad-leaved forest and its main secondary forests (regenerated Lithocarpus forest, Populus bonatii forest, and Alnus nepalensis forest) in Ailao Moutains of Yunnan Province. The results showed that in the primary forest, the carbon storage of woody debris amounted to 36.56 t x hm(-2). Castanopsis wattii, Lithocarpus xylocarpus and L. chintungensis were the main contributors, and most of them were the logs with larger diameter and at intermediate stage of decay. The unique environment of richer precipitation, higher humidity and lower temperature in the study area, and the decay-resistance of hardwood were favorable to the accumulation of woody debris. The three secondary forests had a carbon storage of 1.2-5.0 t x hm(-2), which decreased in the order of regenerated Lithocaropus forest > P. bonatii forest > A. nepalensis forest, showing a tendency of increasing carbon storage with succession course.