Modelling the spatial distribution of beta diversity in Australian subtropical rainforest

Investigation of the spatial distribution of biodiversity among communities or across habitats (beta diversity) is often hampered by a scarcity of biological survey data. This is particularly the case in communities of high floristic diversity, such as the subtropical rainforests of eastern Australia. In contrast, there is excellent spatial coverage of environmental data for this region, such as geology, elevation and climate data. Generalized dissimilarity modelling was used in this study to combine biological survey data and environmental data grids for the investigation and prediction of floristic turnover among vegetation communities at a regional scale. Generalized dissimilarity modelling identified four environmental predictors of floristic turnover in the study region, all of which are linked with moisture stress: radiation of the driest quarter, precipitation of the driest period of the year, slope and aspect. Ten land classes representing largely homogeneous floristics and environment were identified and mapped for the region, allowing significantly greater discrimination than currently available mapping for this region. With increases in evapotranspiration and moisture stress predicted as a result of climate change, these results may allow future floristic shifts to be assessed in relation to regional‐scale gradients in floristic turnover.     

Floristic distributional patterns in a diverse ecotonal area in South America

The Paraguayan territory and region, in the centre of South America, is a huge transition area with a succession of various vegetation types. However, this area has received little attention from researchers, with few works published on its flora and its delimitations. We aimed to identify the most important environmental driving forces and delimit floristic patterns in this region, since understanding the forces that drive floristic variations in this ecotonal region could help comprehend the distribution of vegetation not only in this region but throughout South America. We obtained 1234 tree species occurrence records, 205 geographic coordinates and 23 environmental variables and altitude from the ‘NeoTropTree’ database and verified the influence and contribution of environmental factors through variance partition. We tested the floristic consistency of the different vegetation types using dendrogram, indicator species and ordination analyses. We also constructed multiple linear models to check the correlation between species distribution and environmental variables. We found eight consistent vegetation types. The spatial variables coupled with environmental variables were more important than individual environmental or spatial variables. Among the environmental variables, the aridity index was the most important. Despite the importance of spatial factors, due to environmental heterogeneity, we found a gradient related to climate and edaphic variables related to tree flora. The results confirm that the Paraguayan territory and region can be considered to be a diversified and important ecotone area in South America with respect to tree flora.     

Reproductive phenology of coastal plain Atlantic forest vegetation: comparisons from seashore to foothills

The diversity of tropical forest plant phenology has called the attention of researchers for a long time. We continue investigating the factors that drive phenological diversity on a wide scale, but we are unaware of the variation of plant reproductive phenology at a fine spatial scale despite the high spatial variation in species composition and abundance in tropical rainforests. We addressed fine scale variability by investigating the reproductive phenology of three contiguous vegetations across the Atlantic rainforest coastal plain in Southeastern Brazil. We asked whether the vegetations differed in composition and abundance of species, the microenvironmental conditions and the reproductive phenology, and how their phenology is related to regional and local microenvironmental factors. The study was conducted from September 2007 to August 2009 at three contiguous sites: (1) seashore dominated by scrub vegetation, (2) intermediary covered by restinga forest and (3) foothills covered by restinga pre-montane transitional forest. We conducted the microenvironmental, plant and phenological survey within 30 transects of 25 m × 4 m (10 per site). We detected significant differences in floristic, microenvironment and reproductive phenology among the three vegetations. The microenvironment determines the spatial diversity observed in the structure and composition of the flora, which in turn determines the distinctive flowering and fruiting peaks of each vegetation (phenological diversity). There was an exchange of species providing flowers and fruits across the vegetation complex. We conclude that plant reproductive patterns as described in most phenological studies (without concern about the microenvironmental variation) may conceal the fine scale temporal phenological diversity of highly diverse tropical vegetation. This phenological diversity should be taken into account when generating sensor-derived phenologies and when trying to understand tropical vegetation responses to environmental changes.     

Iteratio: calculating environmental indicator values for species and relevés

Question: Is it possible to translate vegetation maps into reliable thematic maps of site conditions? Method: This paper presents a new method, called Iteratio, by which a coherent spatial overview of specific environmental conditions can be obtained from a comprehensive vegetation survey of a specific area. Iteratio is a database application which calculates environmental indicator values for vegetation samples (relevés) on the basis of known indicator values of a limited number of plant species. The outcome is then linked to a digitalized vegetation map (map of plant communities) which results in a spatial overview of site conditions. Iteratio requires the indicator values of a minimum of 10–20% of the species occurring. The species are given a relative weight according to their amplitudes: species with a narrow range are weighted stronger, species with a broad range are weighted weaker. Conclusion: The method presented here enables a coherent assessment of site conditions on the basis of a vegetation survey and the indicator values of a limited number of plant species.     

Forest classification in an Amazonian rainforest landscape using pteridophytes as indicator species

Habitat classification systems are poorly developed for tropical rainforests, where extremely high plant species richness causes numerous methodological difficulties. We used an indicator species approach to classify primary rainforest vegetation for purposes of comparative wildlife habitat studies. We documented species composition of pteridophytes (ferns and fern allies) in 635 plots (2×100 m) along 8 transects within a continuous rainforest landscape in northeastern Peruvian Amazonia. Considerable floristic variation was found when the data were analyzed using multivariate methods. The obtained forest classification was interpreted with the help of indicator value analysis and known soil preferences of the pteridophyte species. The final classification included four forest types: 1) inundated forests, 2) terrace forests, 3) intermediate tierra firme forests and 4) Pebas Formation forests. This rapid and relatively simple vegetation classification technique offers a practical, quantitative method for large-scale vegetation inventory in complex rainforest landscapes.     

Revealing biogeographical patterns by nonlinear ordinations and derived anisotropic spatial filters

Aim Spatial floristic and faunistic data bases promote the investigation of biogeographical gradients in relation to environmental determinants on regional to continental scales. Our aim was to extract major gradients in the distribution of vascular plant species from a grid‐based inventory (the German FLORKART data base) and relate them to long‐term precipitation and temperature records as well as soil conditions. We present an ordination technique capable of coping with this complex data array. The goal was also to sort out the influence of spatial autocorrelation, assuming floristic autocorrelation is anisotropic. Location Germany, at a spatial resolution of 6′ × 10′. Methods Isometric feature mapping (Isomap) was applied as a nonlinear ordination method. Isomap was coupled to ‘eigenvector‐based filters’ for generating spatial reference models representing spatial autocorrelation. What is novel here is that the derived filters are not based on the assumption of equidirectional autocorrelation. Instead, the so‐called ‘principal coordinates of anisotropic neighbour matrices’ build filters to test the influence of geographical vicinity in directions of high similarity among observations. Results The Isomap ordination of floristic data explained more than 95% of the data variance in six dimensions. The leading two dimensions (representing about 80% of the FLORKART data variance) revealed clear spatial gradients that could be related to independent effects of temperature, precipitation and soil observations. By contrast, the third and higher FLORKART dimensions were dominated by an antagonism of anisotropic spatial autocorrelation and soil conditions. A subsequent cluster analysis of the floristic Isomap coordinates educed the spatial organization of the floristic survey, indicating a considerable sampling bias. Conclusions We showed that Isomap provides a consistent methodical framework for both ordination and derived spatial filters. The technique is useful for tracing the often nonlinear features of species occurrence data to environmental drivers, taking into account anisotropic spatial autocorrelation. We also showed that sampling biases are a conspicuous source of variance in a frequently used floristic data base.     

The inselberg flora of Atlantic Central Africa. I. Determinants of species assemblages

Aims To identify the relative contributions of environmental determinism, dispersal limitation and historical factors in the spatial structure of the floristic data of inselbergs at the local and regional scales, and to test if the extent of species spatial aggregation is related to dispersal abilities. Location Rain forest inselbergs of Equatorial Guinea, northern Gabon and southern Cameroon (western central Africa). Methods We use phytosociological relevés and herbarium collections obtained from 27 inselbergs using a stratified sampling scheme considering six plant formations. Data analysis focused on Rubiaceae, Orchidaceae, Melastomataceae, Poaceae, Commelinaceae, Acanthaceae, Begoniaceae and Pteridophytes. Data were investigated using ordination methods (detrended correspondence analysis, DCA; canonical correspondence analysis, CCA), Sørensen's coefficient of similarity and spatial autocorrelation statistics. Comparisons were made at the local and regional scales using ordinations of life‐form spectra and ordinations of species data. Results At the local scale, the forest‐inselberg ecotone is the main gradient structuring the floristic data. At the regional scale, this is still the main gradient in the ordination of life‐form spectra, but other factors become predominant in analyses of species assemblages. CCA identified three environmental variables explaining a significant part of the variation in floristic data. Spatial autocorrelation analyses showed that both the flora and the environmental factors are spatially autocorrelated: the similarity of species compositions within plant formations decreasing approximately linearly with the logarithm of the spatial distance. The extent of species distribution was correlated with their a priori dispersal abilities as assessed by their diaspore types. Main conclusions At a local scale, species composition is best explained by a continuous cline of edaphic conditions along the forest‐inselberg ecotone, generating a wide array of ecological niches. At a regional scale, these ecological niches are occupied by different species depending on the available local species pool. These subregional species pools probably result from varying environmental conditions, dispersal limitation and the history of past vegetation changes due to climatic fluctuations.     

Land surface phenology as an indicator of biodiversity patterns

With the rapid decline in biodiversity worldwide it is imperative to develop procedures for assessing changes in biodiversity across space. The synoptic view provided by imaging remote sensors constitutes a suitable approach for analyzing biodiversity from local to regional scales. A procedure based on the close relationship between floristic similarity and the similarity in land surface phenology was recently developed and successfully applied to assess diversity patterns using time series imagery acquired by the Moderate Resolution Imaging Spectro-radiometer (MODIS). However, as it depends on high temporal resolution remotely sensed data (e.g., MODIS), the procedure is constrained by the coarse spatial resolution characterizing these high temporal resolution data. Using an optimized technique for image fusion, we combined high temporal resolution data acquired by the MODIS sensor system with moderate spatial resolution data acquired by the Landsat TM/ETM+ sensor systems. Our results show that the MODIS/Landsat data fusion allows the characterization of land surface phenology at higher spatial resolutions, which better corresponded with information acquired within vegetation survey plots established in temperate montane forests located in Wolong Nature Reserve, Sichuan Province, China. As such, the procedure is useful for capturing changes in biodiversity induced by disturbances operating at large spatial scales and constitutes a suitable tool for monitoring and managing biodiversity.     

A numerical analysis of the mesoscale distribution patterns of vascular plants in the subarctic Kevo Nature Reserve, northern Finland

Different numerical techniques were used to detect and describe the major ecological-biogeographical patterns of vascular plant distributions at the meso-scale level in a subarctic region in Finland. The distribution patterns of 231 native taxa in 362 1 km² grid squares of the Kevo Nature Reserve were analysed by two-way indicator species analysis and detrended correspondence analysis, and were subsequently related to twenty-eight geographical, topographical, geological, and vegetational variables using simple discriminant functions and canonical correspondence analysis with associated Monte Carlo permutation tests.
The floristic variation detected reflects variations in environmental factors operative at the regional and local scales. No major broad-scale coherent geographical patterns were detected; instead, the spatial distribution of the grids with a similar floristic composition shows a scattered distribution. All the numerical techniques reveal a major gradient from alpine areas to lowland sites with rivers and rocky outcrops, and the most important explanatory variables for predicting the main floristic variation are all associated with altitude. The floristic patterns represented by the second ordination gradient mainly correlate with the abundance of mires. Partial ordinations indicate that both the geographical and geological variables explain relatively little of the species distributional patterns.
Although the meso-scale approach reveals much about the plant-environment relationships in the study area, the floristic variation appears to be determined mainly by fine-scale factors. In the most heterogeneous grids, the grid size used fails to detect accurately the ecological patterns of the species present.     

Mountain summer farms in Røldal,western Norway –; vegetation classification and patterns in species turnover and richness

This paper discusses vegetation types and diversity patterns in relation to environment and land-use at summer farms, a characteristic cultural landscape in the Norwegian mountains. Floristic data (189 taxa) were collected in 130 4-m² sample plots within 12 summer farms in Røldal, western Norway. The study was designed to sample as fully as possible the range of floristic, environmental, and land-use conditions. Vegetation types delimited by two-way indicator species analysis were consistent with results from earlier phytosociological studies. Detrended correspondence analysis and canonical correspondence analysis show that rather than being distinct vegetation types, the major floristic variation is structured along a spatial gradient from summer farm to the surrounding heathland vegetation. Species richness (alpha diversity) was modelled against environmental variables by generalized linear modelling and compositional turnover (beta diversity) by canonical correspondence analysis. Most environmental factors made significant contributions, but the spatial distance-to-farm gradient was the best predictor of both species richness and turnover. While summer farms reduce mean species richness at the plot scale, the compositional heterogeneity of the upland landscapes is increased, thereby creating ‘ecological room’ for additional vegetation types and species. Within an overall similarity across scales, soil variables (pH, base saturation, LOI, phosphate and nitrogen) differed considerably in their explanatory power for richness and turnover. A difference between ‘productivity limiting’ factors and ‘environmental sieves’ is proposed as an explanation. Species turnover with altitude is relatively low in grasslands as compared to heaths.     

西藏阿里植物群落的间接梯度分析、数量分类与环境解释

根据对西藏阿里地区163个植物群落样地资料进行的多元分析——排序、数量分类与环境解释,给出了该地区植被的基本类型、生态梯度及其与环境因子的定量关系。基本分析方法包括3个步骤：1)通过无倾向对应分析(DCA)的两个排序向量揭示了阿里植被的两个主要生态梯度;2)由该梯度的二维散点图及二元指示种分析(TWINSPAN)分别产生非等级制与等级制的植物群落分类系统;3)以多元回归分析将排序值与环境及地理参数相联系而给出各类型的环境指标——定量环境解释。分析表明,阿里植被类型及其分布主要取决于热量与湿度梯度,前者可通过地理参数,后者则通过土壤特征的数学表达式来定量地确定。两梯度包含的类型、种类与生境差异颇大,由低山暖性荒漠直到高山冰缘植被,从隐带性沼泽与盐生草甸到高原地带性荒漠与草原均各得其位,各有其值。表明该数量分析法对于处理高度生态多样性的植物群落生态信息是十分有效的。     

Partitioning the variation among spatial,temporal and environmental components in a multivariate data set

Abstract We propose a method of partitioning the variation in a multivariate set of data according to (i) environmental variables, (ii) variables describing the spatial structure in the data and (iii) temporal variables. This method is an extension of an existing method for partialling out the spatial component of environmental variation, using canonical analysis. Our proposed method extends this approach by including temporal variables in the analysis. Thus, the partitioning of variation for a data matrix of species’abundances or biomass can include, by our methodology, the following components: (1) pure environmental, (2) pure spatial, (3) pure temporal, (4) pure spatial component of environmental, (5) pure temporal component of environmental, (6) pure combined spatial/temporal component, (7) combined spatial/temporal component of environmental and (8) unexplained. In addition, permutation testing accompanying the analyses allows tests of significance for the relationship between the different components and the species data. We illustrate the method with a set of survey data of penaeid species (prawns) obtained on the far northern Great Barrier Reef, Australia. This extension is a useful tool for multivariate analysis of ecological data from surveys, where space, time and environment commonly overlap and are important influences on observed variation.     

Relationship between topography and the distribution of matorral plant species in the Saharan Atlas: Case of Djebel Amour,Algeria

One of the primary goals in community ecology is to determine the relative importance of processes and mechanisms that control biodiversity. The reduced coverage of vegetation and the severe losses of biodiversity in semiarid matorrals have recently become serious concerns in the Saharan Atlas in southern Algeria. In terms of research, the control of the spatial distribution of vegetation by the topographic factors has not received the attention it merits in Djebel Amour's region. This study combined several multivariate analyses to assess quantitatively the extent and nature of the topographic influence on the distribution of matorral plant species. Thirty plots (20 m × 20 m) were selected according to their physiognomy for the field survey of vegetation and topography in a semiarid mountainous region. The distributions of vegetation differed significantly amongst topographies, indicating different environmental conditions and determine a turnover of important species. Most species colonize downslopes, and only some persist on steep-slopes and summits. The low coverage of vegetation (36.98%) on steep-slopes was due to the serious soil erosion and rockiness 66.5–83.1%. The low average richness in summits (39.67) was due to elevation and negative effect of trees. Elevation, then rockiness and slope were respectively the main topographic factors influencing the distribution of matorral plants. The relationship between topography and distribution could be described by a linear model. Seventeen linear models were built for eleven floristic variables with three topographic factors, elevation, rockiness and slope explained 26–83% of the variance in distribution, also four groups of plant species were listed according to topographic positions. The suitability of plant species to the topographic factors should be considered in the restoration of matorrals in the study area. The results of this study will be useful for selecting the appropriate species and the potential sites for seeding to improving damaged matorral ecosystems in this area. However, additional investigations on soils, water availability and microclimate are required to clarify the mechanism of distribution of plant species in these ecosystems.     

Environmental factors and community dynamics at the southernmost part of the North American Graminetum – I. On the contribution of climatic factors to temporal variation in species composition

Compositional patterns of vegetation and their relationship to temporal and spatial environmental variation, with emphasis on climatic factors, were investigated in plant communities located in the southernmost portion of the North American Graminetum, in central México. Data from 353 samples, obtained in four ecologically contrasting plant communities during 11 years, were analyzed by partial canonical correspondence analysis. Eight climatic variables and eighteen covariables (seventeen edaphic and one resource management) were included in the ordination. A relationship between floristic change and weather variation, once covariables effects were fitted, was examined. Despite a strong contrast in ecological conditions among study sites, a set of four climatic variables was finally found in which each variable contributed independently and statistically (P < 0.01) to the total variance in the vegetation data. Thus, environmental variables other than climatic could not conceal the important role of weather as a mediator of floristic change through time. Summer precipitation and summer maximum temperature showed the highest correlations with the first two species axes, 0.77 and –0.39, respectively. Contribution of these two climatic variables to variance in the vegetation data explained by environmental variables was approximately 81%. Annual species were abundant during rainy years, while abundance of perennial grasses and shrubs showed no clear relationship to weather variation. This study explicitly probes the important role of rain patterns in shaping structure and composition of semiarid communities and considers how other environmental factors can affect plant communities at the southernmost part of the North American Graminetum.     

Do we really need phytosociological classes to calibrate Ellenberg indicator values?

Abstract. Wamelink et al. (2002) calibrated Ellenberg indicator values for acidity and water availability against measured soil pH and measured mean spring groundwater level (MSL), respectively. Linear regression between indicator value and measured value of all the observations gave a poor fit. Regression lines per phytosociological vegetation class, on the other hand, generally described the observations well. In this article we demonstrate that this result is, at least partly, an artefact. First, because the data utilized are likely to contain systematic errors, and second, because a wrong regression model was applied. A sigmoid function for the relation between the indicator value for water availability and MSL gives a far better fit than a linear function does.‘Vegetation class’ is not an obvious choice as an extra explanatory variable for the regression, as it is only a convenient label for vegetation and should not be used as if it were a real independent environmental variable. In general, indicator values of plant species should be calibrated against environmental variables with great care. This implies that researchers should have knowledge about the ecological demands plants make on their environment, as well as about the spatial and temporal variability of this environment.     

Short-term signals of climate change in Italian summit vegetation: observations at two GLORIA sites

Short-term changes occurring in high mountain vegetation were analysed using the data from two Italian sites already part of the GLobal Observation Research Initiative in Alpine environments (GLORIA – central Apennines and southwestern Alps). The study focused on a set of floristic (endemics), structural (life forms) and ecological (thermic vegetation indicator) variables. Vegetation data were collected according to the GLORIA multi-summit standardized method during the last decade. The re-visitation revealed a moderate decrease in regional endemic flora and significant variations in structural and ecological parameters. The increase in caespitose hemicryptophytes in both sites, in suffruticose chamaephytes in the central Apennines and in rosette-forming hemicryptophytes in the southwestern Alps emerged, highlighting the rapid responses of the alpine vegetation to climate warming. The increase in perennial life forms is related with the expansion of graminoids and small woody plants. These life forms seem to be most suitable to face climate warming in Italian summits. The increase in the thermic vegetation indicator exceeds the mean European summits increment, and this is due to the expansion of thermophilic species. Short-term analyses with fine spatial and temporal resolutions are still necessary to improve our understanding concerning species behaviour in high-elevation ecosystems.     

Convergence in the distribution patterns of Europe’s plants and mammals is due to environmental forcing

Aim Our aims were to test: (1) the extent to which vascular plant associations are related in space to mammalian associations, and (2) whether the plant associations are more closely related than the mammalian associations to climate and to a published environmental stratification of Europe. Location Europe, as defined by the following boundaries: 11° W, 32° E, 71° N and 35° N. Methods The analysis is based on presence/absence records of mammal species and plant species with a resolution of 50 km × 50 km. The similarity of the overall spatial structure was tested using a partial Mantel test while controlling for the effect of geographical proximity. To further identify the main spatial components in the datasets, we used k‐means clustering and principal components analysis. The resulting geographical patterns were compared with one another, with climate variables and with the environmental stratification of Europe. Results The clustering of the plant data forms coherent areas that can be interpreted as reflections of floristic regions that are controlled to a large extent by climate and topography. In terms of the correlation between distance matrices, the relationship between plants and mammals is relatively strong. The relationships between mammals and climate, and between plants and climate, are more complex but always statistically significant. There is no evidence that the plant clusters are more closely related than the mammalian clusters to climate, although plant clusters are closer to environmental data than to climate. Main conclusions The clustering patterns of mammals and plants form groups that agree with one another in their spatial extent. The forcing of floristic patterns into coherent entities appears mainly to be caused by climatic variables (temperature, temperature range and rainfall), mediated by elevation differences. The formation of individual plant clusters is also related to species numbers and to local and regional floristic differences. The close correlation between the floristic and faunal patterns suggests that the mammal and plant distributions are controlled by the same environmental variables, although the extent to which the mammals are controlled directly by climate or through the influence of vegetation requires more detailed study.     

Patterns and Determinants of Floristic Variation across Lowland Forests of Bolivia

Floristic variation is high in the Neotropics, but little is known about the factors shaping this variation at the mesoscale. We examined floristic composition and its relationship with environmental factors across 220 1‐ha permanent plots in tropical lowland Bolivia. For each plot, abundance of 100 species (93 tree and 7 palm species ≥10 cm diam) was obtained. Climatic data, related to rainfall seasonality and temperature, were interpolated from all available weather stations in the region, and soil properties, related to texture and fertility, were obtained for each plot. Floristic variation was strongly associated with differences in water availability and temperature, and therefore the climatic gradient shaped floristic variation more strongly than the edaphic gradient. Detrended correspondence analysis ordination divided lowland Bolivia primarily into two major groups (Southern Chiquitano region vs. the Amazon region) and a multiple response permutation procedure distinguished five floristic regions. Overall, the tested environmental variables differed significantly among the five regions. Using indicator species analysis, we distinguished 82 strong indicator species, which had significant environmental preferences for one floristic region. These species can be used as indicators of environmental conditions or to determine which floristic region a certain forest belongs. Given the predicted decreases in rainfall and increases in temperature for tropical lowland forests, our gradient approach suggests that species composition may shift drastically with climate change. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .