Spatial variation of species diversity across scales in an old-growth temperate forest of China

Species richness and abundance are the two most important diversity variables. Species abundance is additive when aggregated across spatial scale, whereas species richness is non-additive. This study analyzes the effect of spatial scale and site on species abundance and richness in a 25-ha temperate forest plot in the Changbai Mountains, northeastern China. The result shows that species abundance and richness are not only dependent on spatial scales, but also dependent on site. Species abundance responds linearly to changes of spatial scale with no intersection in different sites of the study area. However, although species richness also increases with the increase of spatial scale, there are some intersections for the different sites, suggesting that a species-rich site does not always have a high value if the spatial scale is changed. In all, with respect to additive variables, it is relatively easy to extrapolate them from one spatial scale to another spatial scale, as they and the spatial scale usually form a linear relationship. In contrast, non-additive variables are difficult to extrapolate across spatial scales, because they often respond nonlinearly to spatial scale changes. In order to extrapolate these non-additive variables across spatial scales, it is necessary to estimate the relationships between them and spatial scales. As a result, extrapolation of information among spatial scales may be possible, but very difficult, especially for non-additive variables. Because the 25-ha Changbai plot is very small compared to the extent of the world temperate forests, and the vegetation is a relatively uniform type, more such studies in other ecosystems are needed before theories and generalization about scaling effects can be formulated.     

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.     

Relative importance of within-habitat environment,land use and spatial autocorrelations for determining odonate assemblages in rural reservoir ponds in Japan

To clarify the major factors affecting odonate assemblages in rural reservoir ponds among within-habitat environments, land use around ponds and spatial autocorrelation, we surveyed odonate adults (Zygoptera and Anisoptera) in 70 study ponds in Ibaraki Prefecture, Japan, during three sampling periods in 2005. Redundancy analyses (RDA) for these three factor groups were executed to determine their strength in explaining the odonate species composition. Their relative contributions were also evaluated by the method of variation partitioning. A total of 41 odonate species were recorded in the study ponds, and 24 of them, excluding rare species, were used for our analysis. Summed effects including all three factor groups explained approximately 39% of the variation in odonate species composition. We found that spatial autocorrelation was the most important, though the within-habitat environment and land use had comparable effects. We conclude that spatial autocorrelation should be considered in this type of analysis, though we could not clearly explain what caused such a spatial structure. Pond area and debris that had accumulated at the bottom of ponds were selected as the within-habitat environment, and the forests and paddy fields around ponds were selected for land use after the procedure of forward stepwise selection. These results suggest that the recent decrease of forests around the ponds has had a negative effect on the odonate assemblages.     

中国典型森林生态系统乔木层群落物种多样性的空间分布格局及其影响因素

物种多样性地理分布格局及其成因是生物地理学和宏观生态学研究的核心问题之一,基于中国13个典型森林生态系统乔木层群落植物的调查数据,分析物种多样性随经纬度的变化规律,探讨物种多样性空间分布格局的影响因素。结果表明:(1) 13个典型森林生态系统的4个物种多样性指数均随经纬度上升而下降,其中物种丰富度变化更为显著,而Shannon-Wiener指数、Simpson指数和Pielou指数随经度上升变化不显著;(2)相关性分析结果显示,物种多样性指数与植物特性、能量和水分因子的单因素相关关系并不一致。其中,物种丰富度、Shannon-Wiener指数和Simpson指数与年均温、最冷月均温、温度年较差和潜在蒸散量的相关性最显著(P0.01),Pielou指数与年均温、最冷月均温、实际蒸散量、潜在蒸散量和郁闭度有显著相关关系(P0.05);(3)方差分解结果表明,能量和水分的共同作用对物种多样性指数空间分布格局的解释率最高,达到15%—42%;植物特性、能量和水分因子三者共同作用对物种多样性指数空间分布格局解释率次之,为14%—27%;植物特性与能量因子或水分因子两者之间的共同作用以及植物特性和水分因子独立作用对物种多样性指数空间分布格局的解释率较小,其中能量因子对物种多样性指数空间分布格局的单独解释率高于植物特性或水分因子。研究表明能量和水分共同作用是影响大尺度森林乔木层物种多样性空间分布格局形成的主要因素,但植物特性的差异对物种多样性空间分布格局影响也不可忽视。     

Influence of edaphic factors on the spatial structure of inland halophytic communities: a case study in China

Abstract. In order to understand the influence of edaphic factors on the spatial structure of inland halophytic plant communities, a 2.6 km² study site, located on the lower fringe of the alluvial fan of the Hutubi River, in an arid region of China, was sampled and mapped. 105 patches were found to be homogeneous in species composition. Plant species and their coverage were recorded in each patch. 45 patches were randomly selected for the measurement of edaphic variables. A map with quadrat locations and boundaries of patches was digitized into a GIS and related to the vegetation and edaphic data matrices. CCA was used to evaluate the relative importance of edaphic factors in explaining the variation of the species assemblages and to identify the ecological preferences of species. The spatial structure of the communities and the main edaphic factors were analyzed using correlograms, Mantel correlograms and clustering under constraint of spatial contiguity. Gradient analysis showed that there are two distinct vegetation gradients in the study area, one of which is determined mainly by soil moisture (determined by depth to the water table), and the other by soil salinity (determined by electrical conductivity and hydrolytic alkalinity of the first soil layer). However, spatial analyses showed that at the sampling scale the halophytic communities in the study area are structured along one main spatial gradient determined by the water table level. Similar spatial autocorrelation structures between the factors related to the first soil layer and the communities, given our sampling scale, could not be detected. Our results suggest that the relative importance of the effects of different edaphic factors on the spatial structure of halophytic communities is scale-dependent. The partitioning of species variation indicates that in addition to edaphic factors, other factors, such as biotic interactions, may play an important role in structuring these communities.     

Representative estimates of soil and ecosystem respiration in an old beech forest

Respiration has been proposed to be the main determinant of the carbon balance in European forests and is thus essential for our understanding of the carbon cycle. However, the choice of experimental design strongly affects estimates of annual respiration and of the contribution of soil respiration to total ecosystem respiration. In a detailed study of ecosystem and soil respiration fluxes in an old unmanaged deciduous forest in Central Germany over 3 years (2000–2002), we combined soil chamber and eddy covariance measurements to obtain a comprehensive picture of respiration in this forest. The closed portable chambers offered to investigate spatial variability of soil respiration and its controls while the eddy covariance system offered continuous measurements of ecosystem respiration. Over the year, both fluxes were mainly correlated with temperature. However, when soil moisture sank below 23 vol.% in the upper 6 cm, water limitations also became apparent. The temporal resolution of the eddy covariance system revealed that relatively high respiration rates occurred during budbreak due to increased metabolic activity and after leaf fall because of increased decomposition. Spatial variability in soil respiration rates was large and correlated with fine root biomass (r ² = 0.56) resulting in estimates of annual efflux varying across plots from 730 to 1,258 (mean 898) g C m⁻² year⁻¹. Power function calculations showed that achieving a precision in the soil respiration estimate of 20% of the full population mean at a confidence level of 95%, requires about eight sampling locations. Our results can be used as guidelines to improve the representativeness of soil respiration measurements by nested sampling designs, being applied in long-term and large-scale carbon sequestration projects such as FLUXNET and CarboEurope.     

Effects of forest management on epiphytic lichen diversity in Mediterranean forests

Question: What are the responses of epiphytic lichens to the intensity of management along a large environmental gradient in Mediterranean Quercus forests? Location: Central Spain. Methods: This study was carried out on 4590 trees located in 306 forest stands dominated by Quercus faginea or Quercus ilex ssp. ballota. The effect of forest management and other predictor variables on several species diversity indicators were studied. Variables modelled were total species richness, cyanolichen richness and community composition. A large number of predictor variables were included: forest fragmentation (patch size, stand variability), climate and topographic (altitude, slope, sun radiation, annual rainfall and mean annual temperature) and intensity of management. General linear models and constrained ordination techniques were used to model community traits and species composition, respectively. Results: Total richness and especially cyanolichens richness were significantly and negatively affected by the intensity of management. Lichen composition was influenced by management intensity, climatic and topographic variables and stand variability. Conclusions: In Mediterranean forests, human activities related to forestry, agricultural and livestock use cause impoverishment of lichen communities, including the local disappearance of the most demanding species. The conservation of unmanaged forests with a dense canopy is crucial for lichen diversity.     

Effects of experimental N addition on plant diversity in an old‐growth temperate forest

Temperate forest ecosystems have experienced mounting negative effects due to increasing levels of nitrogen (N) deposition. We examined the effects of experimental N addition on plant diversity in an old‐growth temperate forest to test the following hypothesis: Long‐term excessive N addition decreases plant diversity by affecting the growth of plants, which results from changes in the soil nutrient content and a decrease in the soil pH in temperate forests. Experimental N additions were administered at the following levels since 2008: control (0 kg N ha^?1 year^?1), low N (30 kg N ha^?1 year^?1), medium N (60 kg N ha^?1 year^?1), and high N (120 kg N ha^?1 year^?1). Additionally, plant diversity was studied from 2014 to 2016. The results showed that the experimental N additions had significant effects on plant diversity and soil properties in an old‐growth temperate forest. The high‐N treatment decreased the density, cover, and diversity of understory plants, and some herbs even appeared to undergo premature aging, whereas the species diversity of herbs and ferns in the low‐N treatment plots showed a slight increasing tendency. This may have been because the old‐growth temperate forest is an N‐limited ecosystem, so the moderate N input did not show a large influence on plant diversity. However, the long‐term high‐N treatment ultimately reduced plant diversity by changing the soil nutrient contents, decreasing the pH values, and damaging plant growth. Our results suggested that the long‐term excessive N addition negatively affected the forest ecosystem in an N‐limited temperature forest.     

Sap flux of five co-occurring tree species in a temperate broad-leaved forest during seasonal soil drought

In an old-growth forest in Central Germany, sap flux was studied in five broad-leaved tree species that were assumed to differ in drought sensitivity. Under moist soil conditions, average daily sap flux density (J _s) in the outermost xylem varied by a factor of 2.3 among the species (67–152 g cm⁻² per day, n=5 trees per species), and declined in the sequence Fagus sylvatica > Acer pseudoplatanus > Tilia cordata > Carpinus betulus > Fraxinus excelsior. Decreasing soil moisture content (Θ) resulted in linearly reduced J _s in four of the species. During a dry period, J _s was reduced by 44% in T. cordata, 39% in F. sylvatica, 37% in A. pseudoplatanus and 31% in C. betulus compared to sap flux at equal vapour pressure deficit (D) in the wet period. F. excelsior, the only ring-porous species studied, lacked a significant response in J _s to D and Θ. The relative reduction in water use during the dry period was not related to the assumed drought sensitivity of the species as inferred from their abundance in natural woodlands. J _s was positively correlated with tree diameter at breast height (DBH) in three species but decreased with DBH in two species. Dyeing experiments revealed that DBH accounted for 94% of the variation in sapwood area found in a bulk sample of all diffuse-porous trees. This suggests that DBH is a reliable estimator of sapwood area of temperate diffuse-porous species irrespective of species identity. In contrast, sap flux density was found to be greatly dependent on tree species. The estimated whole-plant water use for diffuse-porous trees of a given diameter (49 cm) ranged between 74 and 168 kg per day per species under moist soil conditions. Thus, in temperate mixed forests, species-specific differences in water use can result in a considerable spatial heterogeneity of canopy transpiration.     

Scale-dependent relationships between the spatial distribution of a limiting resource and plant species diversity in an African grassland ecosystem

One cornerstone of ecological theory is that nutrient availability limits the number of species that can inhabit a community. However, the relationship between the spatial distribution of limiting nutrients and species diversity is not well established because there is no single scale appropriate for measuring variation in resource distribution. Instead, the correct scale for analyzing resource variation depends on the range of species sizes within the community. To quantify the relationship between nutrient distribution and plant species diversity, we measured NO₃^- distribution and plant species diversity in 16 paired, modified Whittaker grassland plots in Serengeti National Park, Tanzania. Semivariograms were used to quantify the spatial structure of NO₃^- from scales of 0.4–26 m. Plant species diversity (Shannon-Weiner diversity index; H ) was quantified in 1-m² plots, while plant species richness was measured at multiple spatial scales between 1 and 1,000 m². Small-scale variation in NO₃^- (<0.4 m) was positively correlated with 1-m² H , while 1,000-m² species richness was a log-normal function of average NO₃^- patch size. Nine of the 16 grassland plots had a fractal (self-similar across scales) NO₃^- spatial distribution; of the nine fractal plots, five were adjacent to plots that had a non-fractal distribution of NO₃^-. This finding offered the unique opportunity to test predictions of Ritchie and Olff (1999): when the spatial distribution of limiting resources is fractal, communities should display a left-skewed log-size distribution and a log-normal relationship between net primary production and species richness. These predictions were supported by comparisons of plant size distributions and biomass-richness relationships in paired plots, one with a fractal and one with a non-fractal distribution of NO₃^-. In addition, fractal plots had greater large-scale richness than paired non-fractal plots (1,0–1000 m²), but neither species diversity (H ) nor richness was significantly different at small scales (1 m²). This result is most likely explained by differences in the scale of resource variation among plots: fractal and non-fractal plots had equivalent NO₃^- variation at small scales but differed in NO₃^- variation at large scales (as measured by the fractal dimension). We propose that small-scale variation in NO₃^- is largely due to the direct effects of plants on soil, while patterns of species richness at large scales is controlled by the patch size and fractal dimension of NO₃^- in the landscape. This study provides an important empirical step in understanding the relationship between the spatial distribution of resources and patterns of species diversity across multiple spatial scales.     

长白山三种林型对蛾类群落结构和多样性的影响

选择长白山阔叶红松林、白桦林和落叶松林3种林型,对各林型蛾类的群落结构和多样性进行了比较。结果表明:蛾类各科(亚科)在3种林型中的比例分配明显不同。阔叶红松林中尺蛾科数量占优绝对优势,占47.0%,其次是舟蛾科,为9.3%。优势种为台褥尺蛾,占17.2%;从蛾类总种数和蛾类个体总数上看,白桦林中蛾类物种丰富度、多度和多样性指数明显高于阔叶红松林和落叶松林,优势度指数则最低;白桦林的优势种有尘尺蛾、枞灰尺蛾和双星白枝尺蛾;在落叶松林中,灯蛾科、波纹蛾科和枯叶蛾科的多度都比阔叶红松林和白桦林高,其优势种为阿泊波纹蛾。     

Species and functional group composition of ant communities across an elevational gradient in the Eastern Himalaya

Elevational gradients in mountains show rapid changes in environmental conditions across a small geographic extent. This results in habitat specialization in animal communities which results in changes in species composition across space. We explore changes in species and functional group composition of ants using the first ever data on the distribution of ants across an elevational gradient in the Eastern Himalaya. Ants were sampled from 600 to 2400 m elevations at 200 m intervals using Winklers and pitfall traps. The sampling yielded 166 species of ants from 10,560 individuals, which were then classified into functional groups. We used redundancy analysis to test the effects of environmental factors (temperature, leaflitter volume, understory vegetation) and spatial predictors on species as well as functional group composition of communities at different elevations. Our results show that species diversity within all functional groups decreases towards higher elevations. The functional group composition of ant communities shows a gradient from high evenness at low elevations to being dominated by opportunist species at higher elevations. Redundancy analyses shows that most of the variation in species as well as functional group composition is driven by spatially structured environmental variation. This is most likely due to the high correlation between temperature and elevation. In summary, the changes in species as well as functional group composition are likely driven by a gradient in climate across the elevation gradient.     

Comparison of boosted regression tree and random forest models for mapping topsoil organic carbon concentration in an alpine ecosystem

Soil organic carbon (SOC) plays an important role in soil fertility and carbon sequestration, and a better understanding of the spatial patterns of SOC is essential for soil resource management. In this study, we used boosted regression tree (BRT) and random forest (RF) models to map the distribution of topsoil organic carbon content at the northeastern edge of the Tibetan Plateau in China. A set of 105 soil samples and 12 environmental variables (including topography, climate and vegetation) were analyzed. The performance of the models was evaluated using a 10-fold cross-validation procedure. Maps of the mean values and standard deviations of SOC were generated to illustrate model variability and uncertainty. The results indicate that the BRT and RF models exhibited very similar performance and yielded similar predicted distributions of SOC. The two models explained approximately 70% of the total SOC variability. The BRT and RF models robustly predicted the SOC at low observed SOC values, whereas they underestimated high observed SOC values. This underestimation may have been caused by biased distributions of soil samples in the SOC space. Vegetation-related variables were assigned the highest importance in both models, followed by climate and topography. Both models produced spatial distribution maps of SOC that were closely related to vegetation cover. The SOC content predicted by the BRT model was clearly higher than that of the RF model in areas with greater vegetation cover because the contributions of vegetation-related variables in the two models (65% and 43%, respectively) differed significantly. The predicted SOC content increased from the northwestern to the southeastern part of the study area, average values produced by the BRT and RF models were 27.3 g kg⁻¹ and 26.6 g kg⁻¹, respectively. We conclude that the BRT and RF methods should be calibrated and compared to obtain the best prediction of SOC spatial distribution in similar regions. In addition, vegetation variables, including those obtained from remote sensing imagery, should be taken as the main environmental indicators and explicitly included when generating SOC maps in Alpine environments.     

Vegetation structure and diversity of an ancient temperate deciduous forest in SW Denmark

Bolderslev Skov (113 ha) is the largest contiguous ancient forest remnant in Denmark. The forest has been preserved since 1998 as a strict non-intervention forest reserve. We studied vegetation structure, floristic gradients, and diversity of the forest in 50 plots of 100 m² placed according to a restricted random sampling design. Dominant tree species were Fagus sylvatica, Fraxinus excelsior, Tilia cordata and Quercus robur, which in most parts of the forest form a mixed canopy. Most stands appeared to be of moderate age, 55-80 years old, and large old trees were rare. pH in association with light and thickness of the litter layer were the most important factors in explaining floristic gradients in the forest. Soil moisture (dry to mesic) was not strongly correlated with DCA axes for neither tree nor field layer, but had a significant effect on the distribution of a number of herb species. Forest structure was not important in explaining species distributions. Field layer species richness showed a positive relationship with the pH-gradient. At the scale of 1m² plots we also found a highly significant negative relationship between field layer species richness and the plot-wise Ellenberg indicator value for nitrogen availability. Structure of the tree layer had little effect on field layer species richness. The mixed composition, long continuity, and presence of a high proportion of the regional species pool assigns a high conservation value to Bolderslev Skov and makes it an important site for future studies of the dynamic properties, niche preferences, and inter-specific competition of temperate deciduous forest species. This revised version was published online in June 2006 with corrections to the Cover Date.