Patterns, determinants and models of woody plant diversity in China

What determines large-scale patterns of species richness remains one of the most controversial issues in ecology. Using the distribution maps of 11 405 woody species in China, we compared the effects of habitat heterogeneity, human activities and different aspects of climate, particularly environmental energy, water-energy dynamics and winter frost, and explored how biogeographic affinities (tropical versus temperate) influence richness-climate relationships. We found that the species richness of trees, shrubs, lianas and all woody plants strongly correlated with each other, and more strongly correlated with the species richness of tropical affinity than with that of temperate affinity. The mean temperature of the coldest quarter was the strongest predictor of species richness, and its explanatory power for species richness was significantly higher for tropical affinity than for temperate affinity. These results suggest that the patterns of woody species richness mainly result from the increasing intensity of frost filtering for tropical species from the equator/lowlands towards the poles/highlands, and hence support the freezing-tolerance hypothesis. A model based on these results was developed, which explained 76-85% of species richness variation in China, and reasonably predicted the species richness of woody plants in North America and the Northern Hemisphere.     

Effects of afforestation with pines on woody plant diversity of Mediterranean heathlands in southern Spain

Open heathlands of the Strait of Gibraltar region constitute a singularvegetation type within the Mediterranean Basin owing to their high biodiversitylevels and distinct features in the biological aspects of endemism. However,they have been traditionally depicted as low profitable, tree-less, barrenlandsand have been either overlooked in conservation policies or included inextensive pine-tree afforestation programmes. Nowadays, most of the existingpine plantations are no longer exploited and have become dense pine woodlands.Here we present the results of a comparative analysis aimed to ascertain theeffects of pine stands on the structure and diversity of the woody component ofthese mediterranean heathlands. Ten sampling sites were located within thestudyarea, each consisting of two adjacent subsamples: an open heathland and aneighbouring heathland under pine-tree cover. Woody species richness andoverallshrub cover was much lower in pine-tree heath understoreys. This drop in coverwas more marked for short-lived species (seeders and withering resprouters),implying a decrease of the life-history diversity and also having noticeableconsequences in homogenizing the floristic composition of heath patches underpine trees. Heathland narrow endemics dropped significantly in pine stands,showing greater sensitivity to afforestation than nonendemics. We suggest thatthe open heathlands of the Strait of Gibraltar should be given furtherattentionin EU and IUCN conservation policies. Programmes should be implemented toprogressively eliminate abandoned pine plantations so as to restore andpreservethese highly diverse mediterranean heathlands.     

Spatial heterogeneity in the determinants of woody plant invasion of lowland heath

Questions: 1. What is the scale and extent of spatial variability in factors affecting Betula invasion of heaths? 2. How much effect does each factor have on within‐patch patterns of invasion? 3. How can this understanding aid in managing Betula invasions? Location: Lowland heath of southern England. Methods: Determinants of Betula (both B. pubescens and B. pendula) invasion: biomass density, necromass density, mean vegetation height, P‐availability, soil water content and total Betula seed bank density, were measured at two sites on a 5‐ha sampling grid. Spatial pattern was assessed using geostatistics. Contributions of each determinant to within‐site heterogeneity in predicted Betula seedling densities were estimated by varying variables over their full and interquartile ranges in a statistical model derived from experimental data. Results: Salient spatial trends were revealed: strong autocorrelation over distances of < 50 m for soil factors and more extensive autocorrelation (0 to > 150 m) in vegetation variables and Betula seed bank densities. The latter resulted in single across‐site gradients, the former small, distinct patches. All patterns were overlain with variance that was present at distances of < 17.6 m. Variables displaying spatial pattern also accounted for within‐site heterogeneity in predicted Betula seedling densities but their relative contribution to this varied between sites. Conclusions: Identifiable spatial autocorrelation in factors controlling patch‐scale invasion patterns allows managers to target invasion prone patches, potentially reducing management intensities. Furthermore, management effort may be optimised by spatially de‐coupling Betula seed from safe‐sites. This plan may adaptable to the management of other weeds and open‐land ecosystems.     

Effects of plant biomass, plant diversity, and water content on bacterial communities in soil lysimeters: implications for the determinants of bacterial diversity

Soils may comprise tens of thousands to millions of bacterial species. It is still unclear whether this high level of diversity is governed by functional redundancy or by a multitude of ecological niches. In order to address this question, we analyzed the reproducibility of bacterial community composition after different experimental manipulations. Soil lysimeters were planted with four different types of plant communities, and the water content was adjusted. Group-specific phylogenetic fingerprinting by PCR-denaturing gradient gel electrophoresis revealed clear differences in the composition of Alphaproteobacteria, Betaproteobacteria, Bacteroidetes, Chloroflexi, Planctomycetes, and Verrucomicrobia populations in soils without plants compared to that of populations in planted soils, whereas no influence of plant species composition on bacterial diversity could be discerned. These results indicate that the presence of higher plant species affects the species composition of bacterial groups in a reproducible manner and even outside of the rhizosphere. In contrast, the environmental factors tested did not affect the composition of Acidobacteria, Actinobacteria, Archaea, and Firmicutes populations. One-third (52 out of 160) of the sequence types were found to be specifically and reproducibly associated with the absence or presence of plants. Unexpectedly, this was also true for numerous minor constituents of the soil bacterial assemblage. Subsequently, one of the low-abundance phylotypes (beta10) was selected for studying the interdependence under particular experimental conditions and the underlying causes in more detail. This so-far-uncultured phylotype of the Betaproteobacteria species represented up to 0.18% of all bacterial cells in planted lysimeters compared to 0.017% in unplanted systems. A cultured representative of this phylotype exhibited high physiological flexibility and was capable of utilizing major constituents of root exudates. Our results suggest that the bacterial species composition in soil is determined to a significant extent by abiotic and biotic factors, rather than by mere chance, thereby reflecting a multitude of distinct ecological niches.     

Identifying hotspots of endemic woody seed plant diversity in China

Aim This study aimed to detect distribution patterns and identify diversity hotspots for Chinese endemic woody seed plant species (CEWSPS). Location China. Methods Presence of 6885 CEWSPS throughout China was mapped by taking the Chinese administrative county as the basic spatial analysis unit. The diversity was measured with five indices: endemic richness (ER), weighted endemism (WE), phylogenetic diversity (PD), phylogenetic endemism (PE) and biogeographically weighted evolutionary distinctiveness (BED). Three levels of area (i.e. 1, 5 and 10% of China’s total land area) were used to identify hotspots, but the 5% level was preferred when both the total area of the hotspots identified and the diversity of CEWSPS reached by the hotspots were considered. Results Distribution patterns of CEWSPS calculated with the five indices are consistent with each other over the national extent. However, the hotspots do not show a high degree of consistency among the results derived from the five indices. Those identified with ER and PD are very similar, and so are those with WE and BED. In total, 20 hotspots covering 7.9% of China’s total land area were identified, among which 11 were identified with all the five indices, including the Hengduan Mountains, Xishuangbanna Region, Hainan Island, and eight mountainous areas located in east Chongqing and west Hubei, in east Yunnan and west Guangxi, in north Guangxi, south‐east Guizhou and south‐west Hunan, in north Guangdong and south Hunan, in south‐east Tibet, and in south‐east Hubei and north‐west Jiangxi. Taiwan Island was also identified as a major hotspot with WE, PE and BED. Main conclusions Hotspots of CEWSPS were identified with five indices considering both distributional and phylogenetic information. They cover most of the key areas of biodiversity defined by previous researchers using other approaches. This further verifies the importance of these areas for China’s biodiversity conservation.     

Large mammalian herbivores and woody plant species diversity in Zimbabwe

Study sites from various tree savannas and miombo woodland with adjacentareas with different herbivore species (game and cattle) and stocking rates wereselected from around Zimbabwe. Using the fence-lines separating the adjacentareas, 60 m² plots were placed on both sides of thefence at 18 randomly chosen positions. The identity and number of plants of eachwoody species were recorded in paired 60 m² plots. Therewere no significant differences in the woody species richness between pairedplots for any of the fence-line contrasts. For only two of the fence-linecontrasts were there significant differences (P < 0.05)in the relative abundance of woody species. For the first fence-line contrast,the stocking rates were the same, but the herbivore species present weredifferent, while for the second fence-line contrast, the numbers of herbivorespecies present were the same, but the stocking rates differed. Even thoughthere were no differences in the numbers of woody plant species present, therewere differences in the woody plant species and the numbers of plants of eachspecies: the second most common species was different for three fence-linecontrasts and the third most common species was different for four fence-linecontrasts.     

Effects of herbivores on grassland plant diversity

The role of herbivores in controlling plant species richness is a critical issue in the conservation and management of grassland biodiversity. Numerous field experiments in grassland plant communities show that herbivores often, but not always, increase plant diversity. Recent work suggests that the mechanisms of these effects involve alteration of local colonization of species from regional species pools or local extinction of species, and recent syntheses and models suggest that herbivore effects on plant diversity should vary across environmental gradients of soil fertility and precipitation.     

Influence of landscape complexity and land management on woody plant diversity in northern Spain

Abstract. Changes in land use, particularly the type of forest management, and their spatial pattern, especially woody plant diversity, were related to each other in an area in the Basque region (Spain). The number and diversity of boundaries between landscape patches is related to the development of the spatial complexity of the landscape, C(s), measured in bits and defined as: C(s) = H(b) H(l), where H(b) is the boundary diversity H(1) the spatial heterogeneity generated by these boundaries. As to the vegetation types, changes in land use over the last 50 yr have resulted in a less diversified and more fragmented landscape, especially on a small scale. Forest plantation has become the predominant land use. The frequency of land use changes is studied regarding its influence on the present levels of woody plant diversity.     

中条山中段木本植物群落植物种多样性分析

采用样方法取样,依据重要值指标运用双向指示种分析(TWINSPAN)法,将山西中条山中段植被群系分成16个群丛。应用丰富度指数（R₁、R₂）、Simpson指数（λ）,Shannon-Weiner指数(H′)和均匀度指数(E₁、E₅)研究了16个群丛的植物种多样性。结果表明：群系植物种多样性指数的大小受立地生境和人为活动的综合影响;森林群落的植物种多样性指数明显高于灌丛群落;群落植物种多样性在空间上的差异不仅决定于物种丰富度指数,更与各物种间的均匀度指数密切相关;进而影响不同层次的物种多样性对群落总体物种多样性的贡献。     

The geographical and environmental determinants of genetic diversity for four alpine conifers of the European Alps

Climate is one of the most important drivers of local adaptation in forest tree species. Standing levels of genetic diversity and structure within and among natural populations of forest trees are determined by the interplay between climatic heterogeneity and the balance between selection and gene flow. To investigate this interplay, single nucleotide polymorphisms (SNPs) were genotyped in 24 to 37 populations from four subalpine conifers, Abies alba Mill., Larix decidua Mill., Pinus cembra L. and Pinus mugo Turra, across their natural ranges in the Italian Alps and Apennines. Patterns of population structure were apparent using a Bayesian clustering program, STRUCTURE, which identified three to five genetic groups per species. Geographical correlates with these patterns, however, were only apparent for P. cembra. Multivariate environmental variables [i.e. principal components (PCs)] were subsequently tested for association with SNPs using a Bayesian generalized linear mixed model. The majority of the SNPs, ranging from six in L. decidua to 18 in P. mugo, were associated with PC1, corresponding to winter precipitation and seasonal minimum temperature. In A. alba, four SNPs were associated with PC2, corresponding to the seasonal minimum temperature. Functional annotation of those genes with the orthologs in Arabidopsis revealed several genes involved in abiotic stress response. This study provides a detailed assessment of population structure and its association with environment and geography in four coniferous species in the Italian mountains.     

Effects of prescribed fire on the composition of woody plant communities in southeastern Texas

Abstract. The response to prescribed burning of plant communities ranging from dry to wet habitats was monitored using permanent plots sampled from 1989 to 1993. Temporal controls for fire effects were provided by matched sets of plots protected from fire by newly constructed fire breaks. Changes in species composition were studied by ordination of strata of trees (> 5 cm DBH), small trees (2–5 cm DBH), large saplings (1–2 cm DBH), and small saplings and seedlings (50–140 cm tall). Results show that changes occurred largely in the small tree stratum, in which xeric species increased in importance. Although there were changes in sapling and seedling strata, no clear direction of change was recognized. Fire had little effect on the tree stratum. Of the seven community types under study, three types, sandhill, upland pine, and upperslope pine-oak, were most strongly affected, as indicated by post-fire change in positions of samples representing these communities in ordination space and reduction in understory species abundance. Samples representing the other four mesic and wet communities showed little or no change in their positions. These short-term results indicate that changes in vegetation resulting from fire were small and were mostly restricted to the dry types in which possible compositional change is expected to occur. This differential effect of fire suggests that the influence of fire is secondary to that of topographical and soil gradients in determining vegetation pattern under current fire regimes. Fire seems to reinforce an overall vegetation gradient controlled by soil in southeastern Texas.     

木论喀斯特森林木本植物多样性垂直格局

基于木论国家级自然保护区内西南坡一条垂直样带16个样方的调查资料,运用丰富度指数、物种多样性指数和均匀度指数对木论喀斯特森林木本植物多样性的垂直格局进行研究。结果表明:(1)在16个样方中共记录植株5089个,分属48科90属120种。(2)喀斯特森林木本植物群落具有明显的垂直分布格局,采用等级聚类和非度量多维标度排序方法可将植物群落沿海拔梯度划分为3个类型,分别是:下坡位以小果厚壳桂(Cryptocarya austrokweichouensis)和灰岩棒柄花(Cleidion bracteosum)为主要优势种的群落;中坡位以长管越南茜(Rubovietnamia aristata)和罗伞(Brassaiopsis glomerulata)为主要优势种的群落;上坡位以化香树(Platycarya strobilacea)和密花树(Rapanea neriifolia)为主要优势种的群落。沿海拔梯度升高,水热组合发生变化,地带性植被以及群落内的建群种和优势种都发生了变化。(3)物种生活型方面,常绿树种所占的比例高于落叶阔叶树种,针叶树种仅出现在山顶。(4)沿海拔梯度物种丰富度及α多样性指数中的Shannon-Wiener指数曲线均符合单峰格局,峰值出现在坡腰位置。(5)β多样性指数起伏较大,总体上β多样性随着海拔的升高呈递减的趋势。     

Trait diversity on the phylogeny of cerrado woody species

During the Neogene, the Brazilian cerrado became established as a large‐scale vegetation type. Cerrado lineages started to diversify less than 10 million years ago, coinciding with the rise to dominance of flammable C₄ grasses and the expansion of the savanna biome. Cerrado lineages are strongly associated with adaptations to fire and have sister groups in fire‐free nearby forests, implying that the cerrado formed in situ via adaptive shifts to resist fire. By including phylogeny into the analysis of biological traits, we investigated trait diversity of cerrado woody species in a phylogenetic context, sampling a cerrado site in central Brazil. Decomposing trait diversity along the nodes of a phylogenetic tree of cerrado woody species, we found that the rate of trait diversification was higher in the past, coinciding with the major species diversification of angiosperms in mid‐Cretaceous, long before the cerrado originated. Some more recent adaptive shifts to resist fire, however, must have occurred during the origin and expansion of the cerrado woody flora. Analysing values of each trait separately at the tips of the phylogenetic tree, we found that most trait values were randomly distributed, probably because we analysed only species that had already been filtered by drought, fire, and soil. Analysing values of all traits simultaneously at the tips, we found close to root events and broad, macro‐evolutionary patterns, called ‘global structures’, opposing some lineages, especially Fabaceae and Myrtaceae, with different ecological strategies. Fabaceae presented compound, large, tender leaves, with high nitrogen content due to symbiosis with nitrogen‐fixing bacteria, and Myrtaceae presented simple, small, tough leaves, with low nitrogen and high potassium content. We also found relatively recent events that induced divergence of the evolutionary strategies close to the tips, called ‘local structures’, involving more recent changes in most lineages.     

Effects of above-ground plant species composition and diversity on the diversity of soil-borne microorganisms

A coupling of above-ground plant diversity and below-ground microbial diversity has been implied in studies dedicated to assessing the role of macrophyte diversity on the stability, resilience, and functioning of ecosystems. Indeed, above-ground plant communities have long been assumed to drive below-ground microbial diversity, but to date very little is known as to how plant species composition and diversity influence the community composition of micro-organisms in the soil. We examined this relationship in fields subjected to different above-ground biodiversity treatments and in field experiments designed to examine the influence of plant species on soil-borne microbial communities. Culture-independent strategies were applied to examine the role of wild or native plant species composition on bacterial diversity and community structure in bulk soil and in the rhizosphere. In comparing the influence of Cynoglossum officinale (hound's tongue) and Cirsium vulgare (spear thistle) on soil-borne bacterial communities, detectable differences in microbial community structure were confined to the rhizosphere. The colonisation of the rhizosphere of both plants was highly reproducible, and maintained throughout the growing season. In a separate experiment, effects of plant diversity on bacterial community profiles were also only observed for the rhizosphere. Rhizosphere soil from experimental plots with lower macrophyte diversity showed lower diversity, and bacterial diversity was generally lower in the rhizosphere than in bulk soil. These results demonstrate that the level of coupling between above-ground macrophyte communities and below-ground microbial communities is related to the tightness of the interactions involved. Although plant species composition and community structure appear to have little discernible effect on microbial communities inhabiting bulk soil, clear and reproducible changes in microbial community structure and diversity are observed in the rhizosphere. This revised version was published online in August 2006 with corrections to the Cover Date.     

Landscape structure effects on forest plant diversity at local scale: Exploring the role of spatial extent

Since landscape attributes show different patterns at different spatial extents, it is fundamental to identify how the relation between landscape structure and plant species diversity at local scale varies with scale. Then, it is fundamental to assess the appropriate extent at which landscape factors affect plant species richness at the local scale. To investigate this relation, data on plant species richness of forest communities at plot scale were extracted from a large data set and landscape metrics were calculated around the same plots for a range of extents (250–3000 m). Then, multiple regression models and variance partitioning techniques were applied to assess the amount of variance explained by the landscape metrics on plant species richness for a range of extents. In general, we found that increasing extent of the surrounding landscape analyzed, improved the strength of relationship between the landscape metrics and the properties of plant communities at plot scale. The medium-large extent was most informative as it combined a decent total variance explained with high variance explained by the pure fractions of complexity, fragmentation and disturbance and the minimum of collinearity. In conclusion, we found that it is possible and beneficial to identify a specific extent, where the redundancy in the predictor variables is minimized and the explanatory power of the pure fractions (or single groups) maximized, when examining landscape structure effects on local plant species richness.