Wenyao Liu

Leaf litter decomposition of canopy trees, bamboo and moss in a montane moist evergreen broad-leaved forest on Ailao Mountain, Yunnan, south-west China

Litter decomposition and nutrient release of selected dominant synusiae in an old-growth, evergreen, broad-leaved mossy forest on Ailao Mountain, Yunnan, south-west China, were studied over a 22-month period. The species studied were three dominant tall tree species, Lithocarpus xylocarpus Markg., Lithocarpus chintungensis Hsu et Qian and Castanopsis wattii A. Camus; one dominant understory species (the bamboo Sinarundinaria nitida Nakai); and a mixture of dominant mosses (including Homaliodendron scalpellifolium Fleisch, Symphyodon perrottetti Mont., Herberta longifolissa Steph. and Bazzania albicans Horik.). Fast initial litter decomposition was followed by lower rates. Decomposition rates of canopy species and bamboo leaf litter appear to be controlled by the initial concentration of lignin, nitrogen (N) and phosphorus (P) more than by morphological features of the leaves. The decay rate of moss litter was less correlated with nutrient composition and lignin concentration in initial mass. The order of decomposition rates was Castanopsis wattii > L. xylocarpus > L. chintungensis > bamboo > moss. The decomposition rate constants (k) of the leaf litter for the canopy species L. xylocarpus, L. chintungensis and Castanopsis wattii were 0.62, 0.50 and 0.64, respectively, and 0.40 and 0.22 for bamboo and moss, respectively. Turnover time (1/k) for the three canopy species was 1.61 years, 2.0 years and 1.55 years, respectively, and 2.50 years and 4.55 years for bamboo and moss, respectively. The N and P concentration in the decomposing leaf litter increased in the first 6 months and then decreased over the remaining period. There was a relatively rapid initial loss of potassium (K), followed by a slight increase. Each of calcium (Ca) and magnesium (Mg) decreased with time whereas iron (Fe) and manganese (Mn) increased with time to some extent. Nutrient release from decomposing leaf litter was in the order of K > Mg > Ca > N > P > Mn > Fe, except for bamboo (Sinarundinaria nitida) K > Ca > P > N > Mg > Mn > Fe.

Nutrient fluxes in bulk precipitation, throughfall and stemflow in montane subtropical moist forest on Ailao Mountains in Yunnan, south-west China

A study of nutrient movement through rainfall, throughfall and stemflow, as well as epiphytic bryophyte (moss and liverwort) biomass and its effect on nutrient composition of stemflow, was conducted for 2 y in subtropical montane moist forest (Lithocarpus-Castanopsis association) at Xujiaba, Ailao Mountain National Nature Reserve, Yunnan, south-west China. Base cation and fluxes were increased in throughfall, while NH 4 + -N and NO 3 - -N were reduced relative to precipitation. Annual throughfall inputs of N, P, Ca and S were mainly from precipitation, while most K and two-thirds of Mg throughfall input was due to canopy leaching. Input of Na in the precipitation and throughfall was low in this forest. Net flux (throughfall flux minus bulk precipitation) and deposition ratios (ratio of throughfall flux to bulk precipitation flux) were in the low part of the range reported for other tropical montane rain forests. The throughfall data for this forest reveal generally low cycling rates for mineral elements. Abundant epiphytic bryophytes on bole bark affected the chemical composition of stemflow by selective uptake or release of elements. Relative to trees without epiphytic bryophytes, the annual amounts of total N, NH 4 +-N, Mg, Na and SO 4 2- -S were enhanced, while NO 3 - -N, K, P and Ca were depleted in stemflow. Contributions of N from nitrogen-fixing organisms are likely to be constrained by low temperatures.

Litterfall and nutrient dynamics in a montane moist evergreen broad-leaved forest in Ailao Mountains, SW China

Montane moist evergreen broad-leaved forest, dominated byLithocarpus and Castanopsis species,is the most extensive stand of subtropical mountain in Yunnan Province, SWChina. Litter production, standing crop of litter on forest floor and nutrientreturn patterns were studied over nine years (1991–1999) in a stand ofprimary evergreen broad-leaved forest in northern crest of the Ailao MountainRange. There were significant yearly variations in litter production, which ismainly related with the masting year of canopy species, and exceptionalphysicalevents (strong winds and snow) in the natural forest. The mean annual smalllitterfall is 7.12 t ha−1 yr−1ofwhich leaf litter account for 65% of the total litterfall. The seasonality ofsmall litterfall was bia-modal, with the main one in the late dry season(April–May) and a lesser one in early winter (October–November).Decomposition quotient value was relatively low with 0.58 for total smalllitterfall. Mean large-wood (≥ 2.5 cm in diameter) ranged from0.21 to 1.41 t ha−1 yr−1 with amean of 0.52 t ha−1 yr−1.Concentrations of most elements in leaf and twig were slightly greater in wetmonths than dry months, except for C and K. Woody litter had low N and Pconcentrations compared with the leaf and reproductive parts. Nutrient returntothe soil through small litterfall decrease in the orderC>N>Ca>K>Mg>Mn>Al>P>Fe, while nutrient reserve inlitteron the forest floor was in the declining sequenceC>N>Ca>K>Mg>P>Fe>Al>Mn.

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.

Persistent soil seed banks of the globally significant invasive species, Eupatorium adenophorum, in Yunnan Province, south-western China

Soil cores were collected at different times between the seed germination and dispersal seasons of Eupatorium adenophorum from 19 sites at five stations with different kinds of vegetation in Yunnan, south-western China. Mother plants of E. adenophorum were absent from eight of the sites, and their frequency was low at nine other sites. However, persistent soil seed banks were present at all 19 sites. Seed density in the 0–10 cm soil layer varied from 47 to 13,806 seeds m −2 , and averaged 2199 seeds m −2 . Fifty-seven percent of the seeds of E. adenophorum were in the 0–2 cm soil layer, 24% in the 2–5 cm layer and 19% in the 5–10 cm layer. The percentage of cores from which seedlings emerged ranged from 33–100% across all sites. Seed density and seedling emergence percentages varied significantly among the five stations, and both were positively correlated with abundance of mother plants.

Seasonal variation in density and species richness of soil seed-banks in karst forests and degraded vegetation in central Yunnan, SW China

We studied seasonal variation in density and species richness of seeds in the 0–10 cm soil depth layer in primary, secondary and pine forests, and in shrubland and grassland in the Shilin Stone Forest Geographical Park, Yunnan, SW China. Soil samples were collected four times during the year at 3-month intervals. Seeds from 119 species were identified by germination tests in the soil samples. Density and species richness of seeds of herbaceous plants were greater than woody plants at all five sites throughout the year. Sampling time and site differences had significant effects on the mean number of species and on seed-bank density. Mean number of species per sample increased from February, reached the highest value in May, decreased to the lowest value in August and then increased in November. An exception was in the primary forest, where the highest number of species was found in February. Mean seed-bank density peaked in May at all five sites, and no significant differences were found between densities in February, November and August, except for the primary forest. The peak in seed-bank density in May might be due to dispersal of new seeds of spring-fruiting species, combined with persistence of seeds dispersed in previous years. This seasonal variation of individual species was due primarily to differences in species phenology rather than to differences between sites. Four seasonal seed-bank strategies were identified: two transient and two persistent. At all sites, similar numbers of seeds of herbaceous species were found between seasons, but the number of species of trees and shrubs decreased in August.

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 15N 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.

Adaptation of epiphytic bryophytes in the understorey attributing to the correlations and trade-offs between functional traits

This study explores adaptive strategies of epiphytic bryophytes in the understorey by investigating the photosynthetic characteristics, pigment concentrations and nutrient stoichiometry, as well as other functional traits of three trunk-dwelling bryophytes in a subtropical montane cloud forest in SW China. The results showed that their light-saturated net photosynthetic rate (Anmax−L), light saturation point (Isat), light compensation point (Ic) and dark respiration rate (Rd) were ca 0.55, 106.72, 4.17 and 0.25 μmol m−2 s−1, respectively. Furthermore, the samples demonstrated photosynthetic down-regulation under high irradiance. These photosynthetic characteristics can be explained by higher total chlorophyll concentrations, specific leaf area, chlorophyll per unit leaf N (Chl/N), lower ratio of chlorophyll a to chlorophyll b (Chl a/b) and photosynthetic nitrogen-use efficiency. We suggest that the bryophytes adapted to the shaded understorey microhabitats through a series of correlations and trade-offs between functional traits.

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.

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.