Elements of metacommunity structure of Paraguayan bats: multiple gradients require analysis of multiple ordination axes

Techniques to evaluate elements of metacommunity structure (EMS; coherence, species turnover and range boundary clumping) have been available for several years. Such approaches are capable of determining which idealized pattern of species distribution best describes distributions in a metacommunity. Nonetheless, this approach rarely is employed and such aspects of metacommunity structure remain poorly understood. We expanded an extant method to better investigate metacommunity structure for systems that respond to multiple environmental gradients. We used data obtained from 26 sites throughout Paraguay as a model system to demonstrate application of this methodology. Using presence–absence data for bats, we evaluated coherence, species turnover and boundary clumping to distinguish among six idealized patterns of species distribution. Analyses were conducted for all bats as well as for each of three feeding ensembles (aerial insectivores, frugivores and molossid insectivores). For each group of bats, analyses were conducted separately for primary and secondary axes of ordination as defined by reciprocal averaging. The Paraguayan bat metacommunity evinced Clementsian distributions for primary and secondary ordination axes. Patterns of species distribution for aerial insectivores were dependent on ordination axis, showing Gleasonian distributions when ordinated according to the primary axis and Clementsian distributions when ordinated according to the secondary axis. Distribution patterns for frugivores and molossid insectivores were best described as random. Analysis of metacommunities using multiple ordination axes can provide a more complete picture of environmental variables that mold patterns of species distribution. Moreover, analysis of EMS along defined gradients (e.g., latitude, elevation and depth) or based on alternative ordination techniques may complement insights based on reciprocal averaging because the fundamental questions addressed in analyses are contingent on the ordination technique that is employed.     

稀有种处理对RDA排序结果影响的比较研究

为了解稀有种对RDA排序结果的影响,该研究以北京东灵山华北落叶松林调查数据为例,在RDA排序的基础上,对比分析了未处理稀有种RDA与剔除频度5%、盖度5%的稀有种后RDA排序结果的差异,并用蒙特-卡罗拟合检验分析了二者物种变量和环境变量之间的相关关系,以及用Spearman秩相关系数检验了对应排序轴的相关性。结果表明:(1)蒙特-卡罗拟合检验结果显示未处理稀有种RDA与剔除稀有种RDA各自对应的物种变量和环境变量之间均呈极显著相关关系;(2)从排序轴特征值对物种数据方差以及物种—环境关系解释量来看,剔除稀有种RDA前两排序轴与前四排序轴均有较高的物种-环境关系累积解释量;(3)剔除稀有种前后对应排序轴的Spearman秩相关分析结果表明,尽管未处理稀有种RDA和剔除稀有种RDA在第三轴和第四轴间存在一定的交叉,但二者对应的前四排序轴均呈极显著的一一对应关系(P0.001),相似性极高。总之,结合物种-环境关系的累积解释量及对应排序轴的相关性可知,在环境因子个数较少、研究尺度较小时,使用RDA排序揭示植物种、植物群落和环境因子之间相互作用的生态关系时,剔除稀有种前后RDA排序具有较高吻合性,只是对环境因子的解释趋势稍有差异。     

A comparison of reciprocal averaging and non-centred principal components analysis

Non-centred Principal Components Analysis (NPCA) ordinates sites and species simultaneously, and can be solved either by direct iteration or by eigenvector calculation. The weight of sites and species in the analysis is proportional to their overall abundance. Because of this, the method is not susceptible to distortion by rare species, as is the case with Reciprocal Averaging (RA). Detrending techniques can also be applied to this method to eliminate arch effects.When NPCA was tried with field data, it produced ordination axes that were significantly associated to independently measured environmental variables. In contrast, RA failed to produce axes related to environmental factors, even after the main rare species had been eliminated from the analysis.Abbreviations NPCA Non-centred Principal Components Analysis - RA Reciprocal Averaging     

协惯量分析与典范对应分析在植物群落排序中的应用比较

采用协惯量分析(PCA-CA COIA)和典范对应分析(CCA)两种排序方法, 对北京小龙门林场的黄檗 (Phellodendron amurense)群落进行了分析, 并用Spearman秩相关系数检验了对应排序轴的相关性。两种排序方法得出的结果基本一致, 两者的第一排序轴都反映了海拔高度和坡向对群落分布的影响, 而各自第二、第三排序轴所代表的环境意义有所差异, 并出现了交叉, 但是两者的前3个排序轴均反映了海拔、坡位、土壤厚度和凋落物层厚度的变化趋势, 说明在环境因子个数较少或共线性效应不明显的情况下, 协惯量分析也能达到CCA的分析效果, 并且在排序轴特征值解释量上高于典范对应分析。     

Predicting patterns of vascular plant species richness with composite variables: a meso-scale study in Finnish Lapland

This paper is an attempt, using statistical modelling techniques, to understand the patterns of vascular plant species richness at the poorly studied meso-scale within a relatively unexplored subarctic zone. Species richness is related to floristic-environmental composite variables, using occurrence data of vascular plants and environmental and spatial predictor variables in 362 1 km² grid squares in the Kevo Nature Reserve. Species richness is modelled in two different way. First, by detecting the major floristic-environmental gradients with the ordination procedure of canonical correspondence analysis, and subsequently relating these ordination axes to species richness by generalized linear modelling. Second, species richness is directly related to the composite environmental factors of explanatory variables, using partial least squares regression. The most important explanatory variables, as suggested by both approaches, are relatively similar, and largely reflect the influence of altitude or altitudinally related variables in the models. The most prominent floristic gradient in the data runs from alpine habitats to river valleys, and this gradient is the main source of variation in species richness. Some local environmental variables are also relatively important predictors; the grid squares rich in vascular plant taxa are mainly located in the lowlands of the reserve and are characterised by rivers and brooks, as well as by abundant cliff walls. The two statistical models account for approximately the same amount of variation in the species richness, with more than half of the variation unexplained. Potential reasons for the relatively modest fit are discussed, and the results are compared to the characteristics of the diversity-environment relationships at both broader- and finer-scales.     

Carabid Beetles (Coleoptera,Carabidae) as Indicatorsof Hydrological Site Conditions in Floodplain Grasslands

The relationship of carabid beetle species occurrence patterns and environmental variables characterising the hydrological regime has been studied at the River Elbe in Central Germany. Both flood duration and groundwater depth had major influence on species assemblages as the ordination of study plots mainly followed a gradient along these two variables. The simultaneous ordination of the plots according to species occurrence and environmental parameters showed a highly significant joint structure with the first two axes of a co‐inertia analysis, explaining >98% of the variance. A total of 27 species out of 129 caught fulfilled criteria of fidelity and specificity to the plots of the five clusters revealed by their abiotic conditions and were sufficiently abundant to be suitable indicators for one or a combination of clusters of plots. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Are cloud forest tree structure and environment related in the Venezuelan Andes?

Cloud forest vegetation structure and composition were studied in the Venezuelan Andes at three sites in Mérida State. Although the sites are within 10 to 30 km of each other, climatic, geologic and topographic differences are remarkable. The main purpose of the study was to determine the relationship of specific environmental variables to forest vegetation characteristics, including basal area, tree height, density and diversity, and leaf area index (LAI). At 51 plots, all trees' diameter at breast height >10 cm were recorded and identified. Although the environment at the three sites is distinctive, the tree species composition of the most abundant species was very similar. None of the measured environment variables were significantly correlated with the measured vegetation structure variables, except LAI, which was correlated with slope orientation; LAI showed higher values at south‐facing plots. Tree height was relatively uniform, while basal area was highly variable and reached very high values. Stem densities were in the range reported elsewhere in cloud forests. Multivariate analysis using structure or composition data shows segregation of the plots by site. Principal component analyses by site indicate a minor impact of environmental factors on forest variables. At each site, a particular group of species are correlated with the ordination axes. We conclude that species pools and forest dynamics add to the complexity of the structure of the studied cloud forests.     

Species response curves along environmental gradients. A case study from SE Norwegian swamp forests

Abstract. Vegetation science has relied on untested paradigms relating to the shape of species response curves along environmental gradients. To advance in this field, we used the HOF approach to model response curves for 112 plant species along six environmental gradients and three ecoclines (as represented by DCA ordination axes) in SE Norwegian swamp forests. Response curve properties were summarized in three binary response variables: (1) model unimodal or monotonous (determinate) vs. indeterminate; (2) for determinate models, unimodal vs. monotonous and (3) for unimodal models, skewed vs. symmetric. We used logistic regression to test the influence, singly and jointly, of seven predictor variables on each of three response variables. Predictor variables included gradient type (environmental or ecocline) and length (compositional turnover); species category (vascular plant, moss, Sphagnum or hepatic), species frequency and richness, tolerance (the fraction of the gradient along which the species occurs) and position of species along each gradient. The probability for fitting a determinate model increased as the main occurrence of species approached gradient extremes and with increasing species tolerance and frequency and gradient length. Appearance of unimodal models was favoured by low species tolerance and disfavoured by closeness of species to gradient extremes. Appearance of skewed models was weakly related to predictors but was slightly favoured by species optima near gradient extremes. Contrary to the results of previous studies, species category, gradient type and variation in species richness along gradients did not contribute independently to model prediction. The overall best predictors of response curve shape were position along the gradient (relative to extremes) and tolerance; the latter also expressing gradient length in units of compositional turnover. This helps predicting species responses to gradients from gradient specific species properties. The low proportion of skewed response curves and the large variation of species response curves along all gradients indicate that skewed response curves is a smaller problem for the performance of ordination methods than often claimed. We find no evidence that DCA ordination increases the unimodality, or symmetry, of species response curves more than expected from the higher compositional turnover along ordination axes. Thus ordination axes may be appropriate proxies for ecoclines, applicable for use in species response modelling.     

Gradient analysis of dry grassland vegetation in Denmark

Abstract. We present a gradient analysis of 620 vegetation samples covering most of the floristic and environmental variation in semi‐natural grassland vegetation on well‐drained soils in Denmark. Vegetation was sampled using frequency in subplots. Explanatory variables were surface inclination, aspect, pH, geographical co‐ordinates together with indications of soil type. Detrended Correspondence Analysis revealed four floristic gradients that could be interpreted in ecological terms by measured variables supplemented with site calibrations based on weighted averaging of Ellenberg's indicator values. All four axes were interpreted using rank correlation statistics, and linear and non‐linear multiple regression of sample scores on explanatory variables. The first gradient was from dry calcareous to humid acidic grasslands; the second reflected an underlying gradient in fertility; the third reflected regional differentiation and the fourth was associated with variation in intensity of competition as indicated by association with calibrated Grime‐CSR values for the plots. We applied subset ordination to the data as a supplement to traditional permutation and correlation statistics to assess the consistency of ordination results. DCA axes 1 and 2 were consistent in space and time. This gradient analysis is discussed in a context of plant strategy theory and species diversity models. Ecocline patterns lend support to the view that grazing not only favours the ruderal strategy but also the stress‐tolerant strategy. The low rank of competition as an explanatory variable for the floristical gradients supports the notion that competitive effects play a subordinate role for species composition compared to microclimate and soil conditions in infertile semi‐natural grasslands.     

Butterfly species and traits associated with selectively logged forest in Borneo

Logging can significantly change the structure of rainforest communities. To better understand how logging drives this change, butterflies and environmental variables were assessed within both unlogged and logged forest in Indonesian Borneo. In the whole dataset, we found local environmental variables and geographic distance combined captured 53.1% of the variation in butterfly community composition; 29.6% was associated with measured local environmental variables, 13.6% with geographic distance between sites, and 9.9% with covariation between geographic distance and environmental variables. The primary axis of variation in butterfly community composition represented a disturbance gradient from unlogged to logged forest. Subsequent axes represented gradients influenced by variables such as canopy cover and total tree density. There were significant associations between environmental variables and geographic range and larval host plant use of species. Specifically, butterflies using trees as larval host plants and those with distributions limited to Borneo were more likely to be present in unlogged forest. By contrast, species that tended to be more abundant in logged forest were those with widespread distributions and those using lianas and grasses as larval host plants. The results of this study highlight the importance of environmental variables and disturbance, e.g., selective logging, in structuring rainforest community diversity. Moreover, they confirm how species traits, such as larval food use and geographic distributions can determine patterns of species abundance following environmental change.     

Measuring ecological niche overlap from occurrence and spatial environmental data

Aim Concerns over how global change will influence species distributions, in conjunction with increased emphasis on understanding niche dynamics in evolutionary and community contexts, highlight the growing need for robust methods to quantify niche differences between or within taxa. We propose a statistical framework to describe and compare environmental niches from occurrence and spatial environmental data. Location Europe, North America and South America. Methods The framework applies kernel smoothers to densities of species occurrence in gridded environmental space to calculate metrics of niche overlap and test hypotheses regarding niche conservatism. We use this framework and simulated species with pre‐defined distributions and amounts of niche overlap to evaluate several ordination and species distribution modelling techniques for quantifying niche overlap. We illustrate the approach with data on two well‐studied invasive species. Results We show that niche overlap can be accurately detected with the framework when variables driving the distributions are known. The method is robust to known and previously undocumented biases related to the dependence of species occurrences on the frequency of environmental conditions that occur across geographical space. The use of a kernel smoother makes the process of moving from geographical space to multivariate environmental space independent of both sampling effort and arbitrary choice of resolution in environmental space. However, the use of ordination and species distribution model techniques for selecting, combining and weighting variables on which niche overlap is calculated provide contrasting results. Main conclusions The framework meets the increasing need for robust methods to quantify niche differences. It is appropriate for studying niche differences between species, subspecies or intra‐specific lineages that differ in their geographical distributions. Alternatively, it can be used to measure the degree to which the environmental niche of a species or intra‐specific lineage has changed over time.     

Plant-environment relationships on the Montlake wildlife area,Seattle, Washington,USA

Grassland vegetation on the Montlake fill was analyzed using TWINSPAN. Eight herb communities were recognized. Moisture, proximity to gas vents, and disturbance are the main factors that control species and community distributions. Binary discriminant analysis (BDA) and detrended correspondence analysis (DCA) were used to study species-environment relationships. BDA revealed complex species response patterns and the resultant indicator values were used to interpret the ordination axes. Species distributions are controlled primarily by moisture, but also influenced by soil pH. Multiple regressions revealed little about plant-environment relationships not discovered by BDA. Before robust nonlinear methods are available, BDA, metric ordination with data stratification and nonmetric ordination are methods that can yield satisfactory results in exploratory plant-environment studies. BDA alone is an efficient, useful first approach where response patterns of species are initially unknown.Abbreviations BDA Binary Discriminant Analysis - DCA Detrended Correspondence Analysis     

Environmental Correlates of Tree and Seedling–Sapling Distributions in a Mexican Tropical Dry Forest

Bray and Curtis ordination was used to explore which environmental variables explained importance values and the presence–absence of tropical tree seedlings, saplings and adults in La Escondida-La Cabaña, Sierra de Manantlán, Jalisco, Mexico. The diameters of trees ≥2.5 cm DBH and the presence and height of seedlings and saplings were measured in nine 0.1 ha sites. Four matrices including presence–absence data and importance value indices for trees and seedlings and saplings were analyzed through Bray and Curtis ordination. The matrices were based on density, frequency, and dominance of adult trees as well as seedlings and saplings. The environmental matrix consisted of 18 variables, including elevation, slope, canopy gaps, disturbance, and soil variables. We recorded 63 tree species and 38 seedling and sapling species in the nine sites. The ordination explained 70.9% of the variation in importance value data for trees and 62.6% for seedlings and saplings. The variation explained in presence–absence data for trees was 67.1 and 77.4% for seedlings and saplings. The variance in the ordination axes of seedlings and sapling presence–absence data was poorly explained by the number of gaps in the tree, shrub, or herb layer, suggesting little light specialization by seedlings and saplings. Habitat specialization for soil nutrients appears to be important in explaining the presence–absence of seedlings and saplings. Seedling and sapling specialization along different soil microsites could promote species coexistence in this forest, while heterogeneity in light conditions may instead determine differences in growth and, thus, importance value of trees. We hypothesize that in tropical dry forest in Jalisco, Mexico, a habitat specialization for soil resources is likely more importance at early stages in tree life histories than in later life history.     

Clustering and canonical correspondence analysis of phytoplankton and environmental variables in Swedish lakes

Phytoplankton data consisting of 145 species from a limnological study of lakes from relatively undisturbed areas throughout Sweden were analysed in relation to 11 physical and chemical environmental variables. Three multivariate methods were applied: WPGMA clustering and TWINSPAN for classification, and detrended canonical correspondence analysis (DCCA), a recent technique which extracts ordination axes that can be related directly to variation in the environment. Three types of lakes were recognized consistently: acid humic lakes with Gonyostomum semen as the dominant species, very acid impoverished lakes with rather few, stress-tolerant species, and subarctic lakes with low total biomass but with a varied phytoplankton flora. DCCA allowed a straightforward display of the locations of lakes and species along environmental gradients (including the acidification gradient) reflected in phytoplankton composition. It is suggested that such analyses may be a useful tool for the early detection of environmental change.     

Reducing effort in the use of benthic bioindicators

Hard-bottom invertebrates were visually sampled in a man-modified bay during a harbour enlargement scheme. Several environmental variables were repeatedly measured. Faunal assemblages were analysed using classification and ordination (MDS) techniques at different levels of resolution (species, families and indicators). Indicator taxa were selected from the full species data set by performing preliminary correlations between faunal and environmental variables. Similar results were observed at the three levels investigated. This suggests the existence of redundant information in the species level data set when applied to this case study. The implications of our findings for environmental monitoring are discussed.     

The influence of multi‐scale environmental variables on the distribution of terricolous lichens in a fog desert

Question: How do environmental variables in a hyper‐arid fog desert influence the distribution patterns of terricolous lichens on both macro‐ and micro‐scales? Location: Namib Desert, Namibia. Methods: Sites with varying lichen species cover were sampled for environmental variables on a macro‐scale (elevation, slope degree, aspect, proximity to river channels, and fog deposition) and on a micro‐scale (soil structure and chemistry). Macro‐scale and micro‐scale variables were analysed separately for associations with lichen species cover using constrained ordination (DCCA) and unconstrained ordination (DCA). Explanatory variables that dominated the first two axes of the constrained ordinations were tested against a lichen cover gradient. Results: Elevation and proximity to river channels were the most significant drivers of lichen species cover in the macro‐scale DCCA, but results of the DCA suggest that a considerable percentage of variation in lichen species cover is unexplained by these variables. On a micro‐scale, sediment particle size explained a majority of lichen community variations, followed by soil pH. When both macro and micro‐scale variables were tested along a lichen cover gradient, soil pH was the only variable to show a significant relationship to lichen cover. Conclusion: The findings suggest that landscape variables contribute to variations in lichen species cover, but that stronger links occur between lichen growth and small‐scale variations in soil characteristics, supporting the need for multi‐scale approaches in the management of threatened biological soil crust communities and related ecosystem functions.     

On non-metric multidimensional scaling ordination and interpretation of the matorral vegetation in lowland Murcía

The plant-environment relationships in a range of matorral communities, having different rainfall conditions in semi-arid lowland habitats in Murcía, S.E. Spain, were examined using the non-metric multidimensional scaling ordination technique. Hypotheses on floristic variations were derived based on an interpretative strategy which involved a site configuration rotation, followed by stepwise multiple regression analysis. An environmental data set was used to isolate variables associated with site co-ordinate trends. Results showed that floristic variation in these communities was mainly determined by aspect induced radiation receipt. Besides, most environmental trends in the study areas were found to be oblique to original site ordination axes and configuration rotation seemed to be a prerequisite in quantitative interpretation. The interpretative strategy introduced in this study was effective. It enhanced straightforward, quantitative and objective interpretation whereby inductive inferences on environmental trends could be readily formulated.Abbreviations ARAD radiation receipt surrogate - DEPTH soil depth - DISTURB disturbance index - NMDS non-metric multidimensional scaling - NUT nutrient scalar - SLOPE slope angle - TEXTIN texture index     

Explanations of commonness and rarity in British breeding birds: separating resource use and resource availability

Two of the most prominent explanations for a positive interspecific relationship between local abundance and regional distribution are the resource breadth and the resource availability hypotheses. Here we test these hypotheses by characterising habitat use of British breeding birds using extensive census and environmental data. A group of 85 bird species was considered for study along with 34 land use or environmental variables, which were used to generate four ordination axes by Canonical Correspondence Analysis. Measures of niche breadth and position were derived from these synthetic environmental axes using standard procedures. Across species, none of five measures of abundance and distribution chosen were correlated with niche breadth, whereas four out of five of these measures were correlated negatively with niche position. Repeating the analyses using a method designed to control for phylogenetic non-independence confirmed these general patterns. Birds that tended to use resources that were more atypical of the environment tended to be rarer and thinly distributed, while those using typical resources were common and widely distributed. Performing the analyses on subsets of the data based on species sample sizes did not alter the conclusions. On the assumption that our analyses properly capture patterns of niche use by birds, we find little support for the resource breadth hypothesis, but considerable support for the resource availability hypothesis.     

Biological guidelines for freshwater sediment based on BEnthic Assessment of SedimenT (the BEAST) using a multivariate approach for predicting biological state

Abstract This paper describes the first results for an alternative approach to the development of sediment quality criteria in the nearshore areas of the Laurentian Great Lakes. The approach is derived from methods developed in the United Kingdom for establishing predictive relationships between macroinvertebrate fauna and the physico-chemistry of riverine environments. The technique involves a multivariate statistical approach using (i) data on the structure of benthic invertebrate communities, (ii) functional responses (survival, growth and reproduction) in four sediment toxicity tests (bioassays) with benthic invertebrates; and (iii) selected environmental variables at 96 reference (‘clean’) sites in the nearshore areas of all five Great Lakes (Lakes Superior, Huron, Erie, Ontario and Michigan). Two pattern recognition techniques (using the computer software package PATN) are employed in the analysis: cluster analysis and ordination. The ordination vector scores from the original axes of the pattern analysis are correlated (using CORR in SAS) with environmental variables which are anticipated to be least affected by anthropogenic activities (e. g. alkalinity, depth, silt, sodium etc.). Multiple discriminant analysis (MDA) is used to relate the site groupings from the pattern analysis to the environmental variables and to generate a model that can be used to predict community assemblages and functional responses at new sites with unknown but potential contamination. The predicted community assemblages and functional responses are then compared with the actual benthic communities and responses at a site, and the need for remedial action is determined. The predictive capability of the discriminant model was confirmed by performing several validation runs on subsets of the data. An example of the use of the model for sediment in Collingwood Bay (an area of concern designated by the IJC in Georgian Bay, Lake Huron) is presented and the technique is shown to be more precise in determining the need for remediation than the currently used provincial sediment quality criteria based on Screening Level Concentration (SLC) and laboratory toxicity tests. The ultimate goal of the study is the development of a method to determine the need for, and the success of, remedial action and to predict what benthic communities should look like at a site if it were clean and what responses of organisms in sediment toxicity tests constitute an acceptable end-point.