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.     

Assessing the scale-specific importance of niches and other spatial processes on beta diversity: a case study from a temperate forest

Niche processes and other spatial processes, such as dispersal, may simultaneously control beta diversity, yet their relative importance may shift across spatial and temporal scales. Although disentangling the relative importance of these processes has been a continuing methodological challenge, recent developments in multi-scale spatial and temporal modeling can now help ecologists estimate their scale-specific contributions. Here we present a statistical approach to (1) detect the presence of a space–time interaction on community composition and (2) estimate the scale-specific importance of environmental and spatial factors on beta diversity. To illustrate the applicability of this approach, we use a case study from a temperate forest understory where tree seedling abundances were monitored during a 9-year period at 40 permanent plots. We found no significant space–time interaction on tree seedling composition, which means that the spatial abundance patterns did not vary over the study period. However, for a given year the relative importance of niche processes and other spatial processes was found to be scale-specific. Tree seedling abundances were primarily controlled by a broad-scale environmental gradient, but within the confines of this gradient the finer scale patchiness was largely due to other spatial processes. This case study illustrates that these two sets of processes are not mutually exclusive and can affect abundance patterns in a scale-dependent manner. More importantly, the use of our methodology for future empirical studies should help in the merging of niche and neutral perspectives on beta diversity, an obvious next step for community ecology. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Changes in plant species diversity along a chronosequence of vegetation restoration in the humid evergreen broad-leaved forest in the Rainy Zone of West China

Plant species diversity has been recognized as one of the vital attributes for assessing vegetation restoration. Changes in the diversity may be related to different stages of succession. In this study, 54 sites of humid, evergreen, broad-leaved forest were selected in the Rainy Zone of West China. A chronosequence of the sites was used to study the successive patterns of the diversity in the forest that had undergone natural regeneration for 5 to 350 years and to test the hypothesis that the diversity is maximized in mid-succession. Data were collected simultaneously at different stages of succession, and four α-diversity indices (species richness, Margalef index, Shannon-Wiener index, Pielou Evenness index) and two β-diversity indices (Whittaker index, Sørensen’s index) were calculated for each stratum in each plot. A total of 394 vascular plant species were recorded. From the β-diversity indices, the forest succession may be divided into the early-successional stage (before 50 years), mid-successional stage (from 50 to 300 years), and late-successional stage (after 300 years). In this community, the species diversity and richness were found to be the greatest at the mid-successional stage, followed by the late- and early-successional stages. The results of regression analysis indicated that the richness and Margalef index peaked around the 175th and 165th year, respectively. Shannon-Wiener index values also appeared to follow an approximately humped pattern of succession and were maximal around the 100th year. However, the species evenness did not show any significant relationship with successional age. Our results demonstrate (1) forest restoration is a long-term process and the formation of climax forest requires at least 300 years and (2) the forest has a strong capacity for restoration. Our results also suggest Lindera limprichitii and Machilus pingii as ideal tree species for afforestation because of their wide niche.     

The effects of forest disturbance on diversity of tropical soil nematodes

We provide the first account of the effects of forest disturbance on species richness of nematodes in tropical forest soils, from 24 sites along gradients of disturbance and regeneration in the Mbalmayo Forest Reserve, Cameroon. Species richness was very high. Samples of 200 nematodes from individual soil cores contained a maximum of 89 and an average of 61 species; in total we recorded 431 species and approximately 194 genera. The model of Siemann et al. (1996), predicting that species richness scales as the number of individuals I ^0.5, underestimates nematode diversity 4–6 fold in these samples. Over 90% of specimens cannot be assigned to known species. Although nematode species richness declined with forest disturbance, statistically significant effects were detectable only under the most extreme conditions (active slash-and-burn agriculture and complete mechanical forest clearance) and even here remained at 40% of the richness of near primary sites. Impacts on trophic structure were also small, and there were no significant changes in the maturity index (MI) (Bongers 1990) with disturbance (mean MI across all treatments was very high, at 3.58). In the light of this study, the problems of completing reliable all-taxon inventories in tropical forests are briefly discussed. Received: 22 July 1996 / Accepted: 3 April 1997     

Soil pH and species diversity in coastal dunes

Soil pH was measured at two different spatial scales in coastal dunes on Norderney, North Sea, and in Mecklenburg-Vorpommern, Baltic Sea, Germany. Relationships between the variability in soil pH, species richness and species diversity are presented. Species richness and diversity were highest in grey dunes, where soil pH was at intermediate levels; both variables were lower in yellow and brown dunes. The variability in pH increased with increasing species diversity and also with scale. Overall, soil pH variability decreased with increasing vegetation cover. The lowest pH heterogeneity was found in heath dominated by Empetrum nigrum L. and grey dunes dominated by Campylopus introflexus (Hedw.) Brid. Increasing abundance of dominant species and decreasing species diversity of vegetation apparently reduces soil heterogeneity. Decreasing species diversity of vegetation is likely to explain decreasing variability in soil pH.     

Plant species diversity in abandoned coppice forests in a temperate deciduous forest area of central Japan

We investigated plant species diversity as it related to stand structure and landscape parameters in abandoned coppice forests in a temperate, deciduous forest area of central Japan, where Fagus crenata was originally dominant. The species occurring in the study plots were classified into habitat types based on a statistical analysis of their occurrence bias in particular habitats (e.g., primary forest, coniferous plantation) in the landscape studied. The relationships between stand structure, which reflected the gradient of management, and forest floor plant species diversity (H and J) and richness (number of species per unit area) were not significant. However, these factors did influence the forest floor plant composition of the different types of habitat. According to the multiple regression analysis, species diversity and the richness of forest floor plants was affected by landscape parameters rather than by stand structure. For trees, species richness was mainly affected by the relative dominance of F. crenata, which is one of the stand structure parameters that decreases with intensive management. This is probably because many of the tree species that are characteristic of coppice forests increase after F. crenata have been eliminated by management; these species are not dominant in the original forest, where they are suppressed by F. crenata, the shade-tolerant dominant species. The species diversity (H and J) of trees was positively correlated with some landscape parameters, including the road density around the study plot, which may be associated with the intensity of management activity. The number of disturbance-tolerant species increased with increasing road density. Stand structure mainly affected disturbance-intolerant forest floor plant species and disturbance-tolerant tree species. Thus, the species diversity responses differed between forest floor plants and trees. The impact of forest management on species diversity was more prominent for forest floor plants.     

Tree species diversity and forest structure in relation to microtopography in a tropical freshwater swamp forest in French Guiana

Diversity of tree association and forest structure were analysed in relation to microtopography and flooding intensity in a tropical freshwater swamp forest in the Sinnamary river basin, French Guiana. A 530-m-long vegetation transect was established through a hummock-hollow terrain. Nine 10 m× 50 m sample plots, perpendicular to the main transect, were located so that each was as microtopographically uniform as possible. Trees with dbh (diameter at 1.3 m) 10 cm were censused in all plots and trees with 2 cm dbh < 10 cm in three plots. Sixty tree species belonging to 39 genera and 30 families were recorded. The study area was divided into low and high sites according to microtopography and flooding intensity. According to the Czekanowski similarity matrix, the tree association in low, most frequently flooded, sites differed from that in the high sites under intermediate or low flooding intensity. The low sites had higher stand density and lower species richness than the high sites. Trees with dbh 10 cm in low sites were small and stand basal area (SBA) was about the same in low (69.6 m² ha^–1) and high (64.3 m² ha^–1) sites. The low areas were dominated by Pterocarpus officinalis (38% of stems with dbh 10 cm and 36% of SBA) and Malouetia tamaquarina (26 and 15%). Diospyros guianensis (13.4% of stems with dbh 10 cm and 6.1% of SBA), a Caraipa sp. (14.0 and 7.9%), Lecythis corrugata (6.6 and 3.5%) and emergent Caryocar microcarpum (0.9 and 13.9%) were abundant in high sites. Nodulated legume trees, P. officinalis, Hydrochorea corymbosa and Inga disticha, comprised 44% of stems in the low sites. The abundant nodulation suggests that symbiotic dinitrogen fixation may be an adaptation to N-depleted waterlogged soils. Other adaptive responses were litter accumulation between the buttresses of P. officinalis, which formed hummocks above surface water, and clonal growth habit of M. tamaquarina, which resulted in formation of monospecific groves in low sites.     

Diversity of the understory vascular vegetation in 40 year-old and old-growth forest stands on Vancouver Island,British Columbia

Abstract. We studied plant diversity of the understory vascular vegetation in 40 yr-old plantations (immature stands) and old-growth forest stands on southwestern Vancouver Island, British Columbia, Canada. Site-specific comparisons using several indices of species diversity were made between: (1) immature stands segregated according to the canopy cover and dominant canopy tree species; and (2) immature and old-growth stands. There were no significant differences (P < 0.05) among immature stands in species richness (S) and the Shannon-Wiener index (H′), in relation to the canopy cover or in S, H′ and evenness (E) in relation to the dominant canopy tree species. Using the same indices, the plant diversity varied with edaphic conditions (represented by five site associations) and time (represented by two developmental stages). At both stand- and site levels, plant diversity increased with increasing soil moisture, from slightly dry to moist sites, and with increasing plant-available soil nitrogen in both immature and old-growth stands; and the plant diversity of immature stands across the sites studied was considerably lower than in old-growth stands, regardless of site association. The indices of plant diversity, floristic similarity indices, and species turnover rates indicated that the immature stands had their plant diversity at a minimum, but a drastic loss of diversity expected in the stem exclusion stage had not materialized. We attributed decline in plant diversity to the absence of old-growth structural features in immature stands. Several measures to foster the stand-level diversity were proposed.     

Effects of river level fluctuation on plant species richness, diversity, and distribution in a floodplain forest in Central Amazonia

River levels in Central Amazonia fluctuate up to 14 m annually, with the flooding period ranging from 50 to 270 days between the rising and falling phases. Vast areas of forest along the rivers contain plant species that are well adapted to annual flooding. We studied the effect of flooding level on tree species richness, diversity, density, and composition in lake, river, and stream habitats in Jaú National Park, Brazil. 3051 trees >10 cm diameter (at 1.3 m diameter at breast height, dbh) were measured and identified in 25 10 m × 40 m randomly selected plots in each habitat. Ordination methods and analysis of variance results showed that forested areas near lakes had significantly lower species richness of trees than riverine and streamside habitats. Plot species richness and diversity were strongly negatively correlated with the water level and duration of flooding. The drier (stream) habitat had more total species (54 species of trees) and more unique species of trees (6 tree species) than the riverine (52 tree species; 3 unique species) and lake (33 tree species; 3 unique species) habitats. Species composition overlap among habitats was surprisingly high (42.6–60.6% overlap), almost one-third of the species were found in all three habitat types, and few species were unique to each habitat. We conclude that: (1) duration of flooding has a strong impact on species richness, diversity and plant distribution patterns; (2) most species are adapted to a wide range of habitats and flood durations; and (3) while flood duration may decrease local diversity, it also creates and maintains high landscape-scale diversity by increasing landscape heterogeneity. Received: 20 April 1997 / Accepted: 14 January 1999     

Network properties, species abundance and evolution in a model of evolutionary ecology

We study the evolution of the network properties of a populated network embedded in a genotype space characterized by either a low or a high number of potential links, with particular emphasis on the connectivity and clustering. Evolution produces two distinct types of network. When a specific genotype is only able to influence a few other genotypes, the ecosystem consists of separate non-interacting clusters (i.e. isolated compartments) in genotype space. When different types may influence a large number of other sites, the network becomes one large interconnected cluster. The distribution of interaction strengths--but not the number of connections--changes significantly with time. We find that the species abundance is only realistic for a high level of species connectivity. This suggests that real ecosystems form one interconnected whole in which selection leads to stronger interactions between the different types. Analogies with niche and neutral theory and assembly models are also considered.     

Composition and diversity of the spider fauna in the canopy of a montane forest in Tanzania

Spiders were sampled using insecticide knockdown in an African montane forest in the Uzungwa Mountains of Tanzania. The results are used to discuss the faunal composition at the site and in comparison to other sites, and the implications of the results for estimating spider diversity in Africa are discussed. A total of 5233 adults comprising 149 species were collected from 11 samples covering a total of 906 m² of projected area. Three species contributed 45% of the sample. Previous insecticide knockdown studies of tropical lowland forest canopies have shown a dominance of Theridiidae, Salticidae and Araneidae. In the present study Linyphiidae dominated in abundance and were the second most diverse in terms of species richness. Other abundant families were Oonopidae and Pholcidae, while Theridiidae, Salticidae and Araneidae were rich in species. This supports a previous study, which indicated that the importance of linyphiids increases with altitude. Species richness was predicted using a number of estimators, which produced relatively similar results. Using the abundance-based estimator, Chao 1, the predicted richness for the total area sampled is 183 ± 15 species. This indicates that at least 20% of the area's spider community remains unsampled. A high ratio of undescribed species (approximately 80%) and a relatively high species turnover compared to a site 20 km away within the same forest complex suggests that the number of spiders in Africa could well be much higher than the current, published estimate of 20000 species.     

Spatial processes that maintain biodiversity in plant communities

The composition of communities of sessile organisms, and the change in species diversity with time, is a spatially explicit phenomenon. Three spatial factors clearly affect diversity: (1) the structure and heterogeneity of the landscape that limits species immigration and ultimate community size; (2) neighborhood interactions that determine colonization and extinction rates and influence residence times of local populations; and (3) disturbances that open spatially contiguous areas for recolonization by less abundant species. The importance of these three factors was first reviewed and then examined with a spatially explicit, multi-species model of plant dispersal, competition and establishment, with an assumption of neutrality (all species had equivalent life histories) that reduced the initial dimensionality of the problem. The simulations assumed that the probability of immigration was a linear function of mainland abundance and distance to islands, similar to the equilibrium theory of island biogeography and the unified neutral theory of biodiversity. The rate of increase in species richness was not constant across island sizes, declining as island area became very large. This pattern was explained by the spatial dynamics of colonization and establishment, a non-random process that cannot be explained by passive sampling alone. Simulations showed that population establishment depended critically on rare long-distance dispersal events while population persistence was achieved by the formation of aggregated species distributions that developed through restricted dispersal and local competitive interactions. Nevertheless, species richness always declined to a single species in the absence of disturbances, while up to 40 species could persist to 10,000 years when spatially dependent mortality was added. Further explorations with spatially explicit models will be required to fully appreciate the consequence of land use change and altered disturbance regimes on patterns of species distribution and the maintenance of diversity.     

Seasonal dynamics of arboreal spider diversity in a temperate forest

Measuring and estimating biodiversity patterns is a fundamental task of the scientist working to support conservation and inform management decisions. Most biodiversity studies in temperate regions were often carried out over a very short period of time (e.g., a single season) and it is often-at least tacitly-assumed that these short-term findings are representative of long-term general patterns. However, should the studied biodiversity pattern in fact contain significant temporal dynamics, perhaps leading to contradictory conclusions. Here, we studied the seasonal diversity dynamics of arboreal spider communities dwelling in 216 European beeches (Fagus sylvatica L.) to assess the spider community composition in the following seasons: two cold seasons (I: November 2005-January 2006; II: February-April) and two warm seasons (III: May-July; IV: August-October). We show that the usually measured diversity of the warm season community (IV: 58 estimated species) alone did not deliver a reliable image of the overall diversity present in these trees, and therefore, we recommend it should not be used for sampling protocols aimed at providing a full picture of a forest's biodiversity in the temperate zones. In particular, when the additional samplings of other seasons (I, II, III) were included, the estimated species richness nearly doubled (108). Community I possessed the lowest diversity and evenness due to the harsh winter conditions: this community was comprised of one dominant species together with several species low in abundance. Similarity was lowest (38.6%) between seasonal communities I and III, indicating a significant species turnover due to recolonization, so that community III had the highest diversity. Finally, using nonparametric estimators, we found that further sampling in late winter (February-April) is most needed to complete our inventory. Our study clearly demonstrates that seasonal dynamics of communities should be taken into account when studying biodiversity patterns of spiders, and probably forest arthropods in general.     

Plant community structure,connectance, niche limitation and species guilds within a dune slack grassland

Species phytomass in 40 mm×40 mm quadrats was used to examine community structure in four homogeneous areas of vegetation in sand dune slacks. There were from 9 to 16 common species in the samples, and most parameters of community structure were found to relate consistently to species number. We used covariance between all species at increasing dimensions of aggregations of quadrats to suggest that sociological interaction between species existed at scales of around 200 mm diameter. Generalised variance was of no help in this enquiry. We show that a connectance estimation arrived at from these parameters is, although a real feature of the analysis, apparently unrelated to connectance as understood in the food web literature. There is strong evidence that guild structure, sociological interaction and niche limitation increase with increasing number of species present, so that most diverse assemblage can be said to have a definite community structure.     

Idiosyncratic soil-tree species associations and their relationships with drought in a monodominant Amazon forest

Monodominant forests are characterized by the strong influence of a single species on the structure and diversity of the community. In the tropics, monodominant forests are rare exceptions within the generally highly diverse tropical forest biome. Some studies have shown that tree monodominance may be a transient state caused by successional and demographic variation among species over time. Working in a Brosimum rubescens Taub. (Moraceae) monodominant forest at the southern edge of Amazonia, we tested the hypotheses that local-scale variation in intra- and interspecific spatial patterns of dominant tree species is affected by i) demographic rates of recruitment and mortality following severe droughts, ii) local variation in edaphic properties, and iii) occupation of species in the vertical layer of the forest. We quantified intra- and interspecific spatial patterns and edaphic associations of the five most abundant species using aggregation and association distance indices, and examined changes over time. We found some support for all hypotheses. Thus, intra- and interspecific spatial patterns of most species varied over time, principally after severe drought, emphasizing species-level variability and their interactions in sensitivity to this disturbance, even as B. rubescens monodominance was maintained. While positive and negative spatial associations with edaphic properties provide evidence of habitat specialization, the absence of negative spatial associations of B. rubescens with edaphic properties indicates that this species experiences little environmental restriction, and this may be one of the factors that explain its monodominance. Spatial repulsion and attraction between species in the same and in different vertical layers, respectively, indicates niche overlap and differentiation, while changes over time indicate that the relationships between species are dynamic and affected by drought disturbance.     

Variability of plant species diversity in Swedish natural forest and its relation to atmospheric deposition

Within a subprogram of Integrated Monitoring (IM), understorey vegetation in Swedish natural forests was observed at fifteen reference sites over the country for twelve seasons, 1982–1993. The main task of the subprogram was to assess the impact of atmospheric deposition, mainly sulphur and nitrogen, on natural vegetation through time. The present study is focused on the variability of plant species diversity at community level and the possible impact of sulphur and nitrogen deposition. Species richness, evenness and diversity varied greatly among the sites, and between years within each site. Regarding only coniferous forests the species richness was higher in the north than in the south. But the effects of site condition and atmospheric deposition were not clarified. Changes in species diversity through time differed from site to site. No overall temporal trend was found. The atmospheric deposition of sulphur and nitrogen demonstrated a clear geographical pattern being low in the north-west and high in the south-west. Sulphur deposition declined significantly in Southern Sweden during the period. We concluded that the species diversity of understorey vegetation at the Swedish IM sites was not significantly influenced by atmospheric deposition. The changes observed are explained as natural processes.     

The importance of plot size and the number of sampling seasons on capturing macrofungal species richness

The species-area relationship is an important factor in the study of species diversity, conservation biology, and landscape ecology. A deeper understanding of this relationship is necessary, in order to provide recommendations on how to improve the quality of data collection on macrofungal diversity in different land use systems in future studies, a systematic assessment of methodological parameters, in particular optimal plot sizes. The species-area relationship of macrofungi in tropical and temperate climatic zones and four different land use systems were investigated by determining the macrofungal species richness in plot sizes ranging from 100 m² to 10 000 m² over two sampling seasons. We found that the effect of plot size on recorded species richness significantly differed between land use systems with the exception of monoculture systems. For both climate zones, land use system needs to be considered when determining optimal plot size. Using an optimal plot size was more important than temporal replication (over two sampling seasons) in accurately recording species richness.