Chang-Fu Hsieh

Relations of soil properties to topography and vegetation in a subtropical rain forest in southern Taiwan

Soil chemical properties for a subtropical rain forest in the Nanjenshan Reserve, southern Taiwan, were examined to determine soil-landscape and soil-vegetation relationships. Soil sampling sites were separated into four groups based on landscape features and exposure to the prevailing northeasterly monsoon winds. Corresponding vegetation types were delimited along the first DCA axis. The forest showed a drastic change both in structure and floristic composition along the wind-stress gradient. Redundancy analysis (RDA) showed that both topographic variables and vegetation types were needed to explain the variation in soil data. Soil properties that differed significantly among landforms were pH, available N, CEC, exchangeable Al, K, Ca and Mg. Levels of pH, exchangeable Ca and Mg increased in a downslope direction, and exchangeable Al tended to be higher in the upper slope soils. These trends pointed to the importance of slope processes in redistribution of soil minerals. The main differences in soil properties attributed to the influence of the occupying vegetation were apparently aspect dependent. The contents of available N, exchangeable K, and CEC in the 0–40 cm depth of soils under windward low-stature (mostly sclerophyllous) forest were consistently lower compared to those under the leeward forest. For a given catena, however, soil variability associated with vegetation differences seemed to be confounded by the slope processes.

Scale‐dependent relationships between tree species richness and ecosystem function in forests

1. The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity.

Patterns of plant invasions in China: Taxonomic, biogeographic, climatic approaches and anthropogenic effects

This study was aimed to determine the patterns as well as the effects of biological, anthropogenic, and climatic factors on plant invasions in China. About 270 volumes of national and regional floras were employed to compile a naturalized flora of China. Habit, life form, origin, distribution, and uses of naturalized plants were also analyzed to determine patterns on invasion. Correlations between biological, anthropogenic and climatic parameters were estimated at province and regional scales. Naturalized species represent 1% of the flora of China. Asteraceae, Fabaceae, and Poaceae are the dominant families, but Euphorbiaceae and Cactaceae have the largest ratios of naturalized species to their global numbers. Oenothera, Euphorbia, and Crotalaria were the dominant genera. Around 50% of exotic species were introduced intentionally for medicinal purposes. Most of the naturalized species originated in tropical America, followed by Asia and Europe. Number of naturalized species was significantly correlated to the number of native species/log area. The intensity of plant invasion showed a pattern along climate zones from mesic to xeric, declining with decreasing temperature and precipitation across the nation. Anthropogenic factor, such as distance of transportation, was significantly correlated to plant invasions at a regional scale. Although anthropogenic factors were largely responsible for creating opportunities for exotic species to spread and establish, the local biodiversity and climate factors were the major factors shaping the pattern of plant invasions in China. The warm regions, which are the hot spots of local biodiversity, and relatively developed areas of China, furthermore, require immediate attentions.

Better environmental data may reverse conclusions about niche‐ and dispersal‐based processes in community assembly

Variation partitioning of species composition into components explained by environmental and spatial variables is often used to identify a signature of niche- and dispersal-based processes in community assembly. Such interpretation, however, strongly depends on the quality of the environmental data available. In recent studies conducted in forest dynamics plots, the environment was represented only by readily available topographical variables. Using data from a subtropical broad-leaved dynamics plot in Taiwan, we focus on the question of how would the conclusion about importance of niche- and dispersal-based processes change if soil variables are also included in the analysis. To gain further insight, we introduced multiscale decomposition of a pure spatial component [c] in variation partitioning. Our results indicate that, if only topography is included, dispersal-based processes prevail, while including soil variables reverses this conclusion in favor of niche-based processes. Multiscale decomposition of [c] shows that if only topography was included, broad-scaled spatial variation prevails in [c], indicating that other as yet unmeasured environmental variables can be important. However, after also including soil variables this pattern disappears, increasing importance of meso- and fine-scaled spatial patterns indicative of dispersal processes.

Plant diversity increases with the strength of negative density dependence at the global scale

Maintaining tree diversity Negative interaction among plant species is known as conspecific negative density dependence (CNDD). This ecological pattern is thought to maintain higher species diversity in the tropics. LaManna et al. tested this hypothesis by comparing how tree species diversity changes with the intensity of local biotic interactions in tropical and temperate latitudes (see the Perspective by Comita). Stronger local specialized biotic interactions seem to prevent erosion of biodiversity in tropical forests, not only by limiting populations of common species, but also by strongly stabilizing populations of rare species, which tend to show higher CNDD in the tropics. Science, this issue p. 1389; see also p. 1328 A global analysis of ~3000 species and ~2.4 million trees elucidates variations in tree species diversity between tropical and temperate latitudes. Theory predicts that higher biodiversity in the tropics is maintained by specialized interactions among plants and their natural enemies that result in conspecific negative density dependence (CNDD). By using more than 3000 species and nearly 2.4 million trees across 24 forest plots worldwide, we show that global patterns in tree species diversity reflect not only stronger CNDD at tropical versus temperate latitudes but also a latitudinal shift in the relationship between CNDD and species abundance. CNDD was stronger for rare species at tropical versus temperate latitudes, potentially causing the persistence of greater numbers of rare species in the tropics. Our study reveals fundamental differences in the nature of local-scale biotic interactions that contribute to the maintenance of species diversity across temperate and tropical communities.

Altitudinal zonation of evergreen broad-leaved forest on Mount Lopei, Taiwan

Evergreen broad-leaved forest was studied in a transect on the northwestern slope of Mount Lopei in order to reveal altitudinal zonation in structure and floristic composi- tion and the decisive environmental factors. 20 plots of 20 m x 20 m at altitudes from 540 m to 1320 m were analysed. 144 woody species were found. The results of a DCA ordination clearly pointed to a single dominant altitudinal gradient. Nev- ertheless, wind-exposure associated topography was found to account for additional variation for a given altitudinal range. Along the altitudinal gradient, four dominance-based forest types were recognized. Tree density, species diversity and evenness of the four types differed significantly but total basal area and tree volume were not significantly different. The 95 % turnover range for woody species as measured by the Community Coefficient was calculated as 1030 m, and the 50 % turnover range as 238 m. For the understorey, the change in species composition with altitude was less obvious. Species population structures of 57 sufficiently abundant species re- vealed four characteristic patterns, but most species showed a good fit to the negatively exponential or power function distri- bution and thus appeared to have good reproduction and regular recruitment. Both ANOVA and Redundancy Analysis (RDA) showed that significant differences among forest types were found for most soil variables. Organic C, exchangeable Na and K tended to increase with altitude, while pH and available N showed a reverse trend. There was little evidence that the differences in soil pH and available N were responsi- ble for the variation in forest growth.

The First National Vegetation Inventory in Taiwan

This article describes the successful implementation and preliminary results of the first national vegetation inventory in Taiwan. This project which aimed to establish national criteria for vegetation classification, map island-wide forest vegetation at a scale of 1: 5000 and establish a vegetation data management system was launched in 2003. More than 3000 permanent plots (20 m×20 m) were set up throughout Taiwan, and 792 articles on vegetation of Taiwan were located to acquire vegetation data. Vegetation types were identified according to the physiognomic appearances in aerial photographs. The national vegetation classification system was constructed by referring to field observations, vegetation data, and currently available vegetation classification schemes. This project was finalized in 2008. Up to 67% of native vascular plant species in Taiwan were recorded, and 59% of the total area of Taiwan was mapped in this project. Preliminary vegetation analyses identified 9 major vegetation types and their diagnostic species, constant species and dominant species. The Taiwan Vegetation Information was established and has performed the functions of data management for the current project since 2003. The latest version of the Taiwan Vegetation Classification System was released in 2007. The achievement and experiences of the current project have paved the way for the successful implementation of subsequent large-scale surveys in Taiwan.

Closely-related taxa influence woody species discrimination via DNA barcoding: evidence from global forest dynamics plots.

To determine how well DNA barcodes from the chloroplast region perform in forest dynamics plots (FDPs) from global CTFS-ForestGEO network, we analyzed DNA barcoding sequences of 1277 plant species from a wide phylogenetic range (3 FDPs in tropics, 5 in subtropics and 5 in temperate zone) and compared the rates of species discrimination (RSD). We quantified RSD by two DNA barcode combinations (rbcL + matK and rbcL + matK + trnH-psbA) using a monophyly-based method (GARLI). We defined two indexes of closely-related taxa (Gm/Gt and S/G ratios) and correlated these ratios with RSD. The combination of rbcL + matK averagely discriminated 88.65%, 83.84% and 72.51% at the local, regional and global scales, respectively. An additional locus trnH-psbA increased RSD by 2.87%, 1.49% and 3.58% correspondingly. RSD varied along a latitudinal gradient and were negatively correlated with ratios of closely-related taxa. Successes of species discrimination generally depend on scales in global FDPs. We suggested that the combination of rbcL + matK + trnH-psbA is currently applicable for DNA barcoding-based phylogenetic studies on forest communities.

Classification of the High-Mountain Coniferous Forests in Taiwan

Vegetation of boreal coniferous forests has been extensively studied in many areas of northern Eurasia and North America, but similar forests in the high mountains of subtropical and tropical eastern Asia have been poorly documented so far. This paper, focusing on such forests, is the first phytosociological study at a national scale in Taiwan. The relevés from the National Vegetation Diversity Inventory and Mapping Project database were used to define vegetation types of the high-mountain coniferous forests and to characterize their distribution in Taiwan. Environmental variables such as aspect, elevation, soil rockiness and slope were related to species composition. Cluster analysis was used to classify vegetation plots and establish groups that were interpreted as nine associations belonging to two alliances. The alliance Juniperion squamatae represents woodlands and forests scattered in the subalpine belt, in which Juniperus squamata dominates the canopy and subalpine meadow species occur in the understorey. The Abieti kawakamii-Tsugion formosanae alliance includes forests dominated by Abies kawakamii and Tsuga chinensis var. formosana with shade-tolerant herb species in the upper montane belt. In addition to regional vegetation description, an identification key for the studied forests was developed based on the classification tree technique.

Local neighborhood communities in the understory play a critical role by affecting regeneration niches and subsequent community assembly in a montane cloud forest

Regeneration niches for tree seedlings are defined as possessing the biotic and abiotic requirements to successfully replace adults. However, two critical discrepancies obscure our understanding of the role of regeneration niches in forest community assembly: a weak association between seedling assemblages and environmental factors, and an uncoupling between seedling and adult-tree assemblages. However, understory plants, which may interact with microenvironments and seedlings, may be a better spatial descriptor of regeneration habitats. To test this, the spatial variations of seedling assemblages in a montane cloud forest of Taiwan were analyzed in terms of their association with neighborhood assemblages of understory plants, the shrub layer, and overstory trees, as well as environmental variables. We found that the understory plant spatial structure contributed most in explaining spatial variations of the seedling assemblages (especially for widespread Lauraceae and patchy Fagaceae) and facilitated the identification of patches of different regeneration habitats for specific seedling assemblages. Moreover, among these regeneration patches, tall herbs affected seedling density and diversity differently. We found segregation between tall herbs and Lauraceae seedlings, indicating that tall herbs shape seedling assemblages and uncouple the association between seedling and adult stages. However, positive covariations between seedlings/tall herbs and between seedlings/adults were found for Fagaceae and Pentaphylacaceae in different regeneration patches, suggesting that positive, neighboring effects and dispersal limitation may cause the patchy distribution of seedling assemblages and affect their coupling with adults. Thus, the understory plant spatial structure shapes seedling assemblages and provides a better link to spatial associations with regeneration habitats.