Spatial pattern of diversity in an old-growth temperate forest in Northeastern China

Species diversity has attracted particular attention because of its significance for helping determine present species performance and likely future community composition. The spatial pattern of species diversity (species richness, abundance and Shannon diversity) in Changbai temperate forest in Northeastern China was studied to investigate the present and likely causes for the formation of spatial patterns. To fulfill this goal, three aspects of diversity were addressed: 1) changes in the relationships of the diversity variables, species richness, abundance and Shannon diversity, to sampling area and sampling design. The three diversity variables were found to respond to sampling area in a dissimilar way. Sampling design had no significant effect on the diversity variable-area curves. The power function, which was derived under the assumption that the forest was in equilibrium, did not fit the observed species-area curves, indicating that the Changbai temperate forest was probably not in equilibrium. 2) Variograms, used to examine the spatial structure of species diversity, showed that the spatial structure of species diversity in the Changbai temperate forest was weakly anisotropic. 3) Partitioning the variation of species diversity into spatial and environmental factors indicated that the spatial pattern of the Changbai forest community was unpredictable, probably because there were many undetermined processes controlling its development.     

Plant Community Structure in Tropical Rain Forest Fragments of the Western Ghats,India1

Changes in tree, liana, and understory plant diversity and community composition in five tropical rain forest fragments varying in area (18–2600 ha) and disturbance levels were studied on the Valparai plateau, Western Ghats. Systematic sampling using small quadrats (totaling 4 ha for trees and lianas, 0.16 ha for understory plants) enumerated 312 species in 103 families: 1968 trees (144 species), 2250 lianas (60 species), and 6123 understory plants (108 species). Tree species density, stem density, and basal area were higher in the three larger (> 100 ha) rain forest fragments but were negatively correlated with disturbance scores rather than area per se. Liana species density, stem density, and basal area were higher in moderately disturbed and lower in heavily disturbed fragments than in the three larger fragments. Understory species density was highest in the highly disturbed 18‐ha fragment, due to weedy invasive species occurring with rain forest plants. Nonmetric multidimensional scaling and Mantel tests revealed significant and similar patterns of floristic variation suggesting similar effects of disturbance on community compositional change for the three life‐forms. The five fragments encompassed substantial plant diversity in the regional landscape, harbored at least 70 endemic species (3.21% of the endemic flora of the Western Ghats–Sri Lanka biodiversity hotspot), and supported many endemic and threatened animals. The study indicates the significant conservation value of rain forest fragments in the Western Ghats, signals the need to protect them from further disturbances, and provides useful benchmarks for restoration and monitoring efforts.     

海南岛霸王岭垂直带热带植被物种多样性的空间分析

以常用的Shannon-Wiener指数,Simpson指数以及新近提出的两个非参数指数(Chao多样性指数和二阶刀切法多样性指数),计测了海南岛霸王岭垂直带上不同植被类型(包括热带低山雨林、热带山地雨林、热带云雾林、热带山地矮林)的物种多样性.在此基础上,借助地理信息系统的空间分析技术,模拟绘制了整个梯度的多样性变化谱.总体而言,样带下部的物种多样性高于上部,或者说热带低山雨林与热带山地雨林的物种多样性高于热带云雾林与热带山地矮林,而南部稍高于北部.这种基于插值方法的空间分析技术为探索植被结构分析提供了一条新的途径.     

海南岛吊罗山山地雨林物种多样性

在海南岛吊罗山热带山地雨林设置两５０００ｍ＾２的样地,分别从物种多样性指数,种－多度曲线,空间格局多样性,时间格局多样性,组成格局多样性对其进行了研究。结果表明海南岛吊罗山山地雨林物种多样性指数Ｓｈａｎｎｏｎ－Ｗｅａｖｅｒ和Ｓｉｍｐｓｏｎ指数分别为４．１９５和０．０２１,均匀度为０．８８０和２．９６２。     

Dynamics of Forest Landscape Patterns Along the Main Road in Xishuangbanna

In this paper, we investigated dynamics of forest landscape along the main roads in Xishuangbanna from 1976 to 1988, and from 1988 to 2003, using FRAGSTATS software and the GIS technology, based on the results interpreted from the Landsat MSS TM ETM imageries in 1976, 1988, 2003. Some representative quantitative indices of landscape diversity, including mean patch area, percent of landscape , perimeter-area fractal dimension , aggregation , Shannon′s diversity index , and Shannon′s equality index were used to describe the changes in the spatial pattern of landscape elements. Comparing with whole region of Xishuangbanna during last 27 years , the results showed that the man-made landscape (rubber plantation) and non-forested landscape along the main road expanded faster and their percentage were greater, whereas the natural landscape ( primitive forest patches) reduced faster and its percentage was smaller . There were obviouseffects of road on landscape which the forest coverage was decreased , and rubber plantation landscape was increased with the distance from the road . The natural forest landscape had been fragmented. Especially, the tropical seasonal rain forest was affected severely, while the area and contagion of rubber plantation and shrub increased. The whole landscape pattern along the main road tended to be diversiform, equal and fragmented. The landscape was still dominated by natural landscapes but this dominance was weakening . A case study about the displacement of landscape patch centroids showed that the spatial centroids of many types of landscape, including rubber plantation, tropical seasonal rain forest, mountain rain forest, shrub and non-forested land had moved away from the road. All of these changes in landscape pattern would lead the deterioration of these region environments.     

The species-area relationship in the hoverfly (Diptera, Syrphidae) communities of forest fragments in southern France

The effect of forest fragmentation was studied in hoverfly communities of 54 isolated forests (0.14–171 ha) in south west France. The positive relationship between species richness and wood patch area was investigated by testing the three hypotheses usually put forward to explain it: 1) the sampling effect hypothesis, 2) the patch heterogeneity hypothesis, 3) the hypothesis of equilibrium between distance from other patch (colonisation) and surface area of the patch (extinction). The syrphid species were divided into 3 ecological groups, based on larval biology as summarized in the "Syrph the Net" database: non forest species, facultative forest species and forest species. A total of 3317 adults belonging to 100 species, were captured in the 86 Malaise traps. Eight species were non forest (N=16), 65 facultative forest (N=2803) and 27 forest species (N=498).
Comparison of the slopes of the species-area curves for species richness and species density per forest patch showed a strong sampling effect in the species-area relationship. Wood patch heterogeneity increased with wood patch area and positively influenced hoverflies richness. Less isolated wood patches presented high richness of forest species and low richness of non forest species. Only forest species richness seemed to respond to the equilibrium between surface area and isolation. Depending on which hypothesis explained best the species-area relationship, management recommendations to mitigate fragmentation effects were formulated at various spatial scales and for different stakeholders.     

Biodiversity assessment in incomplete inventories: leaf litter ant communities in several types of Bornean rain forest

Biodiversity assessment of tropical taxa is hampered by their tremendous richness, which leads to large numbers of singletons and incomplete inventories in survey studies. Species estimators can be used for assessment of alpha diversity, but calculation of beta diversity is hampered by pseudo-turnover of species in undersampled plots. To assess the impact of unseen species, we investigated different methods, including an unbiased estimator of Shannon beta diversity that was compared to biased calculations. We studied alpha and beta diversity of a diverse ground ant assemblage from the Southeast Asian island of Borneo in different types of tropical forest: diperocarp forest, alluvial forest, limestone forest and heath forests. Forests varied in plant composition, geology, flooding regimes and other environmental parameters. We tested whether forest types differed in species composition and if species turnover was a function of the distance between plots at different spatial scales. As pseudo-turnover may bias beta diversity we hypothesized a large effect of unseen species reducing beta diversity. We sampled 206 ant species (25% singletons) from ten subfamilies and 55 genera. Diversity partitioning among the four forest types revealed that whereas alpha species richness and alpha Shannon diversity were significantly smaller than expected, beta-diversity for both measurements was significantly higher than expected by chance. This result was confirmed when we used the unbiased estimation of Shannon diversity: while alpha diversity was much higher, beta diversity differed only slightly from biased calculations. Beta diversity as measured with the Chao-Sørensen or Morisita-Horn Index correlated with distance between transects and between sample points, indicating a distance decay of similarity between communities. We conclude that habitat heterogeneity has a high influence on ant diversity and species turnover in tropical sites and that unseen species may have only little impact on calculation of Shannon beta diversity when sampling effort has been high.     

西双版纳热带雨林树种多样性的尺度效应

基于西双版纳20hm2森林动态监测样地内直径≥1cm的树种资料,分析了该样地树种香农多样性指数和辛普森多样性指数及其方差和变异系数在7个取样尺度(5m×5m、10m×10m、20m×20m、25m×25m、50m×50m、100m×100m、200m×250m)的变化规律,结果表明:(1)香农指数-面积曲线在尺度100m×100m有一个峰值,而辛普森指数-面积曲线在尺度为20m×20m有一个峰值。(2)香农指数和辛普森指数的方差随尺度的增加而减小,其中香农指数的方差在尺度100m×100m上最小,辛普森指数在尺度20m×20m方差最小,表明香农多样性指数在100m×100m的尺度上所获得的多样性信息比较可靠,而辛普森指数在20m×20m的尺度上所获得信息比较可靠。(3)多样性指数的方差和变异系数远大于随机模型拟合的数值,说明样地内树种不是随机分布。     

Diversity and community composition of ectomycorrhizal fungi in a dry deciduous dipterocarp forest in Thailand

Large forest areas of South-East Asia, are dominated by the Dipterocarpaceae tree family, which contains many important timber species. Unlike many other tropical trees, Dipterocarpaceae rely on ectomycorrhizal (ECM) root symbiosis for their mineral nutrition. This study aims to document the richness and community composition of ECM fungi in a dry deciduous forest in Thailand. Combining morphological and molecular identification methods revealed 69 species of ECM fungi that belong to 17 phylogenetic lineages. The /russula-lactarius, /tomentella-thelephora, /sordariales, /sebacina and /cantharellus lineages were the most species-rich. The fungal richness is comparable to other tropical rain forest sites, but the phylogenetic community structure has elements of both tropical and temperate ecosystems. Unlike tropical rain forests, the Cenococcum geophilum complex was one of the most frequent fungal taxa that had a relatively high ITS genetic diversity over the small sampling area. This study provides the first snapshot insight into the fungal community of dry dipterocarp forests. However, it is necessary to broaden the spatial and temporal scales of sampling to improve our understanding of the below-ground relations of dry and humid tropical forests.     

Forest Modification Affects Diversity (But Not Dynamics) of Speciose Tropical Pyraloid Moth Communities

We studied temporal and spatial dynamics of extremely diverse moth ensembles (Lepidoptera: Pyraloidea) along a gradient of forest disturbance ranging from undisturbed primary tropical rain forest to different kinds of modified forest and open cultivated land at the margin of Mount Kinabalu National Park (Sabah, East Malaysia). We sampled moths by light trapping during two periods (March‐May and August‐September 1997). We collected a total of 7724 individuals representing 680 species during 78 light‐trapping nights at six study sites. Species diversity (Fisher's α) of ensembles in undisturbed primary forest was distinctly higher than in disturbed or secondary forest. More pyraloid moths were attracted in undisturbed primary forest. Samples from disturbed primary or old‐growth secondary forest were statistically indistinguishable from the undisturbed primary forest ensemble in regard to species composition. Thus, pyraloid ensembles from disturbed forest with tall trees remaining appeared to represent impoverished subsets of the undisturbed primary forest community. The more heavily disturbed sites had a distinct fauna and showed a stronger faunal differentiation among each other. Four species of the genus Eoophyla, in which aquatic larvae feed on algae in fast‐running streams benefited prominently from forest disturbance. Temporal variation of ensembles was remarkably concordant across the disturbance gradient. Relative abundance variation of the commonest species was identical at all sites. Overall, pyraloid moths responded more sensitively to anthropogenic habitat alteration than most other moth taxa studied thus far in tropical regions and allowed for an analysis of diversity patterns at a high temporal resolution.     

Spatial Variation in Army Ant Swarm Raiding and its Potential Effect on Biodiversity

Invertebrate communities of the tropical rain forest floor are highly diverse, characterized by patchy species distribution patterns and high variation in species density. Spatial variation in the foraging activity of swarm raiding army ants, prime invertebrate predators in tropical rain forests, is discussed as a mechanism contributing to these patterns, but highly resolved long‐term data on army ant raiding on the local and landscape scale are hitherto lacking. In this study, 196 positions in 11 study sites in a tropical rain forest in western Kenya were continuously monitored over ~4 mo for the occurrence of swarm raids of army ants. Using population simulation analyses, the consequences of army ant raiding for prey communities were assessed. We found an unexpectedly high variation in raid rates at the study site and landscape scale. The weekly chance of communities to become raided by army ants was on average 0.11, but ranged from 0 to 0.50 among the 196 positions. Simulating population developments of two Lotka–Volterra species—showing slight trade‐offs between competitive strength and resistance to army ant raids—in the real raiding landscapes showed that the observed spatial variation in raid rates may produce high prey diversity at larger spatial scales (due to high β‐diversity) and strong variation in species density. Our results indicate that high spatial variation in army ant swarm raiding is a mechanism capable of generating patchy species distribution patterns and maintaining the high biodiversity of invertebrate communities of the tropical rain forest floor.     

Additive partitioning of rarefaction curves and species-area relationships: unifying alpha-, beta- and gamma-diversity with sample size and habitat area

Additive partitioning of species diversity is widely applicable to different kinds of sampling regimes at multiple spatial and temporal scales. In additive partitioning, the diversity within and among samples ( α and β ) is expressed in the same units of species richness, thus allowing direct comparison of α and β . Despite its broad applicability, there are few demonstrated linkages between additive partitioning and other approaches to analysing diversity. Here, we establish several connections between diversity partitions and patterns of habitat occupancy, rarefaction, and species–area relationships. We show that observed partitions of species richness are equivalent to sample-based rarefaction curves, and expected partitions from randomization tests are approximately equivalent to individual-based rarefaction. Additive partitions can also be applied to species–area relationships to determine the relative contributions of factors influencing the β -diversity among habitat fragments.     

粤北南方红豆杉植物群落的物种多样性和种群格局

在用样方法取样的基础上 ,分析粤北南方红豆杉生长地植物群落的物种多样性和种群分布格局 ,结果表明 :在调查地南方红豆杉出现于 5种类型的群落中 ;从物种多样性指数的平均值来看 ,灌木层的物种多样性大于乔木层 ,但乔木层和灌木层之间的 Shannon指数和群落均匀度无明显的差异 ;从乔木层和灌木层的变异系数看 ,丰富度、Shannon指数和群落均匀度均表现为乔木层 >灌木层。所调查的南方红豆杉生长地的植物群落的多样性指数和均匀度高于其他类群的南亚热带常绿阔叶林 ,但低于海南岛的热带雨林类群。南方红豆杉生长地植物群落中各优势种群均呈集群分布 ,且集群强度也都较高。南方红豆杉种群在大多数样地中亦呈集群分布     

Don’t miss the forest for the trees! Evidence for vertical differences in the response of plant diversity to disturbance in a tropical rain forest

Ecological studies in tropical rain forests traditionally focus on trees above a threshold diameter at breast height (dbh), since ignoring plant species of the other structural compartments is believed to be an acceptable tradeoff between exhaustiveness and effectiveness. However, the consequences of missing species below a threshold dbh value have been largely neglected so far. We evaluated whether the response of species diversity of ≥10-cm dbh trees was similar to the response of other structural ensembles (namely treelets, saplings, and terricolous herbs) in a lowland tropical rain forest, to three disturbance regimes: natural gap dynamics (control), and selective logging with and without additional thinning. We studied forest vegetation composition and diversity in a 20-yr replicated field experiment comprising nine 1 ha permanent plots established in a semi-deciduous rain forest of the Congo Basin and equally distributed among the three treatments. Once corrected by stem density, species richness was similar between logged (20 years since logging) and untouched old-growth forest stands with respect of trees, but higher with respect of treelets. As disturbance intensity increased, species richness increased within sapling layers but decreased within herb layers, while species spatial turnover (beta diversity) increased in both cases. Regarding the parameters of the partitioned rarefaction curves and relative abundance distribution curves, no correlation was found between trees and any of the other structural compartments. Whilst tree and treelet species composition was similar among treatments, the understories still reflected past disturbance intensity, with a strong response of the sapling and herb layers. These results show that ecological studies based solely on tree layers (dbh ≥ 10 cm) are misleading because their response to disturbance cannot be used as a surrogate for the response of other structural ensembles. Long-lasting effects of anthropogenic disturbance on the sapling bank and the herb layer may durably influence the long-term forest dynamics. Since overstory but not understory plant communities have recovered from human disturbances 20 years after silvicultural operations, African tropical rain forest ecosystems may not be as resilient to selective logging as previously thought.     

滇南热带雨林物种多样性的取样面积探讨

通过不同面积样方种数的比较、物种多样性指数等的计算和逐步扩大样地面积的调查, 本文研究了滇南热带雨林种数/面积关系、个体/种数关系、物种多样性及树种的频度分布规律, 认为滇南热带雨林群落学调查的最适取样(样方)面积是0.25 hm², 这个面积接近该类型热带雨林理论上的群落最小表现面积。 为能体现一个具体森林(群落)类型的基本的植物区系组成, 需设置4～5个这样的样方(总面积累计1 hm²以上)。从与世界不同地区热带雨林的比较亦可见,云南热带雨林有类似的树种频度分布规律,单位面积上的物种多样性比典型的东南亚低地热带雨林稍低,但比非洲的热带雨林要高。     

Soil Effects on Forest Structure and Diversity in a Moist and a Dry Tropical Forest

Soil characteristics are important drivers of variation in wet tropical forest structure and diversity, but few studies have evaluated these relationships in drier forest types. Using tree and soil data from 48 and 32 1 ha plots, respectively, in a Bolivian moist and dry forest, we asked how soil conditions affect forest structure and diversity within each of the two forest types. After correcting for spatial effects, soil‐vegetation relationships differed between the dry and the moist forest, being strongest in the dry forest. Furthermore, we hypothesized that soil nutrients would play a more important role in the moist forest than in the dry forest because vegetation in the moist forest is less constrained by water availability and thus can show its full potential response to soil fertility. However, contrary to our expectations, we found that soil fertility explained a larger number of forest variables in the dry forest (50 percent) than in the moist forest (17 percent). Shannon diversity declined with soil fertility at both sites, probably because the most dominant, shade‐tolerant species strongly increased in abundance as soil fertility increased.     

Inferring species abundance distribution across spatial scales

A long-standing problem in ecology is to understand how the species–abundance distribution (SAD) varies with sampling scale. The problem was first characterized by Preston as the veil line problem. Although theoretical and empirical studies have now shown the nonexistence of the veil line, this problem has generated much interest in scaling biodiversity patterns. However, research on scaling SAD has so far exclusively focused on the relationship between the SAD in a smaller sampling area and a known SAD assumed for a larger area. An unsolved challenge is how one may predict species–abundance distribution in a large area from that of a smaller area. Although upscaling biodiversity patterns (e.g. species–area curve) is a major focus of macroecological research, upscaling of SAD across scale is, with few exceptions, ignored in the literature. Methods that directly predict SAD in a larger area from that of a smaller area have just started being developed. Here we propose a Bayesian method that directly answers this question. Examples using empirical data from tropical forests of Malaysia and Panama are employed to demonstrate the use of the method and to examine its performance with increasing sampling area. The results indicate that only 10-15% of the total census area is needed to adequately predict species abundance distribution of a region. In addition to species abundance distributions, the method also predicts well the regional species richness.     

Detecting successional changes in tropical forest structure using GatorEye drone-borne lidar

Drone-based remote sensing is a promising new technology that combines the benefits of ground-based and satellite-derived forest monitoring by collecting fine-scale data over relatively large areas in a cost-effective manner. Here, we explore the potential of the GatorEye drone-lidar system to monitor tropical forest succession by canopy structural attributes including canopy height, spatial heterogeneity, gap fraction, leaf area density (LAD) vertical distribution, canopy Shannon index (an index of LAD), leaf area index (LAI), and understory LAI. We focus on these variables’ relationship to aboveground biomass (AGB) stocks and species diversity. In the Caribbean lowlands of northeastern Costa Rica, we analyze nine tropical forests stands (seven second-growth and two old-growth). Stands were relatively homogenous in terms of canopy height and spatial heterogeneity, but not in their gap fraction. Neither species density nor tree community Shannon diversity index was significantly correlated with the canopy Shannon index. Canopy height, LAI, and AGB did not show a clear pattern as a function of forest age. However, gap fraction and spatial heterogeneity increased with forest age, whereas understory LAI decreased with forest age. Canopy height was strongly correlated with AGB. The heterogeneous mosaic created by successional forest patches across human-managed tropical landscapes can now be better characterized. Drone-lidar systems offer the opportunity to improve assessment of forest recovery and develop general mechanistic carbon sequestration models that can be rapidly deployed to specific sites, an essential step for monitoring progress within the UN Decade on Ecosystem Restoration.     

西双版纳勐宋轮歇演替区鸟类多样性及食果鸟研究

结合退耕还林、生态保护和山区经济持续发展的需要,在西双版纳勐宋轮歇休闲自然演替林地、传统保护使林地和原始自然林地等3类8种生境中,采用样方法并辅以挂网捕鸟法,研究了鸟类的多样性、食果鸟类及其与森林动态的关系。结果表明：①随着轮歇休闲时间递增和植被层次增加,鸟类多样性增加,鸟类群落逐渐复杂化;②鸟类多样性H、Hmax和J等参数变化情况在干季和雨季类似;③食果传播植物种子的鸟类沟通了各林地之间的联系,它们的活动对植被的演替和扩展至关重要。讨论分析认为,鸟类多样性与生境植被结构相互作用,协同发展;在退耕还林工作中要重视食果类及其生态功能的作用,注意保护原始森林和鸟类物种,利用和促进食果鸟类的自然生态功能。