Accounting for tree spatial distribution in a comparison of plot sizes and shapes in dense forest and woodland in Benin (West Africa)

The study examined simultaneously, the effect of tree spatial distribution, inventory plot size and shape on the estimation error of basal area in two contrasting environments. Twenty and fifteen square plots of 1 ha each (divided into 100 quadrats of 0.01 ha) were randomly set in dense forest and woodland, respectively. Thirteen subplots of various shapes and sizes were obtained from the association of adjacent quadrats. Estimation error was calculated using residual mean square of one‐way ANOVA, based on replications of subplot within 1 ha plots. Tree spatial distribution was measured using Green index. Weighted linear regression and mixed effect models were applied to Box & Cox transformed data. In general, the estimation error of basal area decreased with increase in subplot size. However, the effects of tree spatial distribution and plot shape varied with the vegetation type. Where trees tended to be aggregated, estimation error increased with degree of aggregation, and rectangular plots of 0.24 ha produced an acceptable precision. It was concluded that 0.24 ha rectangular plots can be used in tropical environments where the target parameters vary constantly according to one direction, while square plots of the same size are optimal for reliable analysis in case of randomness.     

On tree intensity estimation for forest inventories: Some statistical issues

The effect of the plot shape, number of subplots and their spatial arrangement on the sample variance for spatially explicit point populations is analysed for a simple intensity estimator. We derive the sample variance and covariance for sampling designs involving more than one subplot. Some numerical approximations are also presented. If a clustered point pattern has to be sampled, the best strategy to reduce the sample variance is to consider as many rectangular subplots as possible, for a prescribed total sample area, distributed over a grid. In contrast, if a regular point pattern is to be sampled, then a single circular subplot should be considered. If we assume that the point configuration is Poisson, then we can consider any subplot shape and spatial distribution ensuring no overlapping between the subplots. A case study in forestry is considered to assess the validity of our results.     

Plot shape effects on plant species diversity measurements

Abstract. Question: Do rectangular sample plots record more plant species than square plots as suggested by both empirical and theoretical studies? Location: Grasslands, shrublands and forests in the Mediterranean‐climate region of California, USA. Methods: We compared three 0.1‐ha sampling designs that differed in the shape and dispersion of 1‐m² and 100‐m² nested subplots. We duplicated an earlier study that compared the Whittaker sample design, which had square clustered subplots, with the modified Whittaker design, which had dispersed rectangular subplots. To sort out effects of dispersion from shape we used a third design that overlaid square subplots on the modified Whittaker design. Also, using data from published studies we extracted species richness values for 400‐m² subplots that were either square or 1:4 rectangles partially overlaid on each other from desert scrub in high and low rainfall years, chaparral, sage scrub, oak savanna and coniferous forests with and without fire. Results: We found that earlier empirical reports of more than 30% greater richness with rectangles were due to the confusion of shape effects with spatial effects, coupled with the use of cumulative number of species as the metric for comparison. Average species richness was not significantly different between square and 1:4 rectangular sample plots at either 1‐ or 100‐m². Pairwise comparisons showed no significant difference between square and rectangular samples in all but one vegetation type, and that one exhibited significantly greater richness with squares. Our three intensive study sites appear to exhibit some level of self‐similarity at the scale of 400 m², but, contrary to theoretical expectations, we could not detect plot shape effects on species richness at this scale. Conclusions: At the 0.1‐ha scale or lower there is no evidence that plot shape has predictable effects on number of species recorded from sample plots. We hypothesize that for the mediterranean‐climate vegetation types studied here, the primary reason that 1:4 rectangles do not sample greater species richness than squares is because species turnover varies along complex environmental gradients that are both parallel and perpendicular to the long axis of rectangular plots. Reports in the literature of much greater species richness recorded for highly elongated rectangular strips than for squares of the same area are not likely to be fair comparisons because of the dramatically different periphery/area ratio, which includes a much greater proportion of species that are using both above and below‐ground niche space outside the sample area.     

Comparison of dwarf bamboos (Indocalamus sp.) leaf parameters to determine relationship between spatial density of plants and total leaf area per plant

The relationship between spatial density and size of plants is an important topic in plant ecology. The self‐thinning rule suggests a ?3/2 power between average biomass and density or a ?1/2 power between stand yield and density. However, the self‐thinning rule based on total leaf area per plant and density of plants has been neglected presumably because of the lack of a method that can accurately estimate the total leaf area per plant. We aimed to find the relationship between spatial density of plants and total leaf area per plant. We also attempted to provide a novel model for accurately describing the leaf shape of bamboos. We proposed a simplified Gielis equation with only two parameters to describe the leaf shape of bamboos one model parameter represented the overall ratio of leaf width to leaf length. Using this method, we compared some leaf parameters (leaf shape, number of leaves per plant, ratio of total leaf weight to aboveground weight per plant, and total leaf area per plant) of four bamboo species of genus Indocalamus Nakai (I. pedalis (Keng) P.C. Keng, I. pumilus Q.H. Dai and C.F. Keng, I. barbatus McClure, and I. victorialis P.C. Keng). We also explored the possible correlation between spatial density and total leaf area per plant using log‐linear regression. We found that the simplified Gielis equation fit the leaf shape of four bamboo species very well. Although all these four species belonged to the same genus, there were still significant differences in leaf shape. Significant differences also existed in leaf area per plant, ratio of leaf weight to aboveground weight per plant, and leaf length. In addition, we found that the total leaf area per plant decreased with increased spatial density. Therefore, we directly demonstrated the self‐thinning rule to improve light interception.     

Spatial pattern analysis in ecology based on Ripley's K-function: Introduction and methods of edge correction

Abstract. Spatial pattern analysis based on Ripley's K-function is a second-order analysis of point patterns in a twodimensional space. The method is increasingly used in studies of spatial distribution patterns of plant communities, but the statistical methods involved are sometimes poorly understood or have been modified without evaluating the effects on results. The procedures of field data acquisition, statistical analysis, and the test for the null hypothesis of complete spatial randomness are described and the presentation of results is discussed. Different methods of edge correction were tested on a computer-generated random pattern and a mapped distribution of a Mediterranean shrubland. The inclusion of buffer zones around mapped plots describes the spatial pattern most accurately, but may not warrant the additional labour involved. Three variations of the weighted edge correction yielded comparable results for the distribution patterns tested. The toroidal edge correction may give biased results for non-random patterns. Recommendations for standardisation of the statistical procedures and data presentation are given.     

Shape matters in sampling plant diversity: Evidence from the field

The identification of shape and size of sampling units that maximises the number of plant species recorded in multiscale sampling designs has major implications in conservation planning and monitoring actions. In this paper we tested the effect of three sampling shapes (rectangles, squared, and randomly shaped sampling units) on the number of recorded species. We used a large dataset derived from the network of protected areas in the Siena Province, Italy. This dataset is composed of plant species occurrence data recorded from 604 plots (10 m × 10 m), each divided in a grid of 16 contiguous subplot units (2.5 m × 2.5 m). Moreover, we evaluated the effect of plot orientation along the main environmental gradient, to examine how the selection of plot orientation (when elongated plots are used) influences the number of species collected. In total, 1041 plant species were recorded from the study plots. A significantly higher species richness was recorded by the random arrangement of 4 subplots within each plot in comparison to the ‘rectangle’ and ‘square’ shapes. Although the rectangular shape captured a significant larger number of species than squared ones, plot orientation along the main environmental gradient did not show a systematic effect on the number of recorded species. We concluded that the choice of whether or not using elongated (rectangular) versus squared plots should dependent upon the objectives of the specific survey with squared plots being more suitable for assessing species composition of more homogeneous vegetation units and rectangular plots being more suited for recording more species in the pooled sample of a large area.     

The effects of habitat edges and trampling on understorey vegetation in urban forests in Helsinki,Finland

Question: How do habitat edges and trampling affect mesic (Myrtillus type) understorey vegetation in fragmented urban forests, and what are the relative strengths and spatial extent of these effects? Location: The city of Helsinki, southern Finland. Methods: Vegetation sample plots were placed at different distances from the forest edge. At each distance we selected plots on, next to and away from paths of different levels of wear, and the covers and frequencies of plant species were sampled. We used generalized linear mixed models and ordination techniques to test our hypotheses. Results: The effect of the edge penetrated up to 50 m into the forest interior. Only light effects of trampling were found in seemingly untrampled areas, in addition to the direct effects of trampling on the paths. Both trampling and edge effects changed the forest understorey species composition. Trampling changed plant species composition locally on paths, and provided opportunities for new species to establish in previously unbroken forest vegetation. Urban forest edges were characterised by species better adapted to sunny, warm and dry conditions. These species, such as grasses, replaced more sensitive forest species, such as dwarf shrubs and mosses. We observed a large number of light demanding deciduous trees at the edges, which may be one of the factors increasing soil fertility and consequently promoting the high abundance of fast growing species, such as grasses and herbs related to more fertile site types. Conclusions: We recommend that urban forest fragments left within urban development should be at least 2–3 ha in size and that the shape of the patch and the number of potential recreational users should be taken into account when preserving true forest understorey vegetation from the effects of edges and trampling.     

Multi-scale comparisons of cliff vegetation in Colorado

We developed a nested vegetation sampling protocol to sample the plant diversity on south-facing cliffs and cliff bases in Jefferson County, Colorado. The multi-scale plots included three nested spatial scales, 1 m², 20 m², and 40 m². We compared plant species richness and species diversity among large cliffs, medium cliffs, small cliffs, and non-cliff sites using Hill's diversity numbers (N ₀, N ₁, and N ₂) for the 1-m² quadrats. Species richness (N ₀) was calculated for the 20-m² and 40-m² plots. Our results indicate that plant species diversity on the cliff faces did not increase with increasing cliff area. This pattern was consistent at all three sampling scales. A model selection was run to determine if plant species diversity values on the cliff faces were associated with cliff variables. None of the cliff variables measured were good predictors of diversity at the 1-m² scale. However, at the 20-m² scale, canyon differences and a positive relationship with increasing cliff surface roughness explained 70% of the variability in species richness. Although most plant species sampled on the cliff faces were also found in the base plots, 13 species were sampled only on the cliff faces. Additionally, dry south facing cliffs support a mix of xeric and mesic plants indicating that cliffs may provide unique microenvironments for plant establishment. This revised version was published online in August 2006 with corrections to the Cover Date.     

Disentangling environmental and spatial processes of community assembly in tropical forests from local to regional scales

Understanding patterns and mechanisms of variation in the compositional structure of communities across spatial scales is one of the fundamental challenges in ecology and biogeography. In this study, we evaluated the effects of spatial extent (i.e. size of study region) on: 1) whether community composition can be better explained by environmental (i.e. niche‐based) or spatial (e.g. dispersal‐based) processes ; and 2) how climate and soils contribute to the influence of environment on plant community composition. We surveyed community composition across a network of 398 forest plots spanning a ～4000 m elevational gradient in the Madidi region in northwestern Bolivia. Using redundancy analyses and hierarchical variation partitioning, we disentangled the effects of environmental and spatial predictors on species composition, further decomposing the environmental effect between its climatic and soil components. We repeated analyses for 200 sub‐regions ranging in spatial extent from ～250 to ～17 500 km². Our analyses show a high degree of idiosyncrasy in results that come from different sub‐regions. Despite this variability, we were able to identify various important patterns in the structure of tropical plant communities in our study system. First, even though sub‐regions varied in size by nearly two orders of magnitude, the total amount of explained variation in community composition was scale independent; at all spatial scales, environment and space accounted for about 25% of the differences in community composition among plots. Second, the measured environmental effect was higher than the spatial effect on average and in the vast majority of sub‐regions. This was true regardless of the spatial extent of analysis. Finally, we found that both climatic and soil variables accounted for significant fractions of variation, but climate was always more important than soils.     

Cross-scale drivers of plant trait distributions in a fragmented forest landscape

During community assembly, plant functional traits are under selective pressure from processes operating at multiple spatial scales. However, in fragmented landscapes, there is little understanding of the relative importance of local-, patch- and landscape-scale processes in shaping trait distributions. Here, we investigate cross-scale influences of landscape change on traits that dictate plant life history strategies in re-assembling plant communities in a fragmented landscape in eastern China. Using forest dynamics plots (FDPs) on 29 land-bridge islands in which all woody plants have been georeferenced and identified to species, we characterized and derived two composite measures of trait variation, representing variation across the leaf economics spectrum and plant size. We then tested for trait shifts in response to local-, patch- and landscape-scale factors, and their potential cross-scale interactions. We found substantial community-wide trait changes along local-scale gradients (i.e. forest edge to interior): more acquisitive leaf economic traits and larger sized species occurred at edges, with a significant increase in trait means and trait range. Moreover, there were significant cross-scale interaction effects of patch and landscape variables on local-scale edge effects. Altered spatial arrangement of habitat in the surrounding landscape (i.e. declining habitat amount and increasing patch density), as well as decreasing area at the patch level, exacerbated edge effects on traits distributions. We suggest that synergistic interactions of landscape- and patch-scale processes, such as dispersal limitation, on local-scale environmental filtering at edges, together shape the spatial distributions of plant life history strategies in fragmented plant communities.     

Aquatic plant community composition and distribution along an inundation gradient at two ecologically-distinct sites in the Pantanal region of Brazil

In spite of its size and biological significance, we know little about the ecology of the Pantanal, a 140,000 km² floodplain in west-central Brazil. Increasing human pressures make this lack of understanding particularly critical. Using transects and 1 m² circular plots, we documented floristic composition and interacting-environmental conditions associated with littoral herbaceous vegetation along inundation gradients at two ecologically-distinct sites in the Pantanal. We recorded water depth and percent cover for each species in Baía Piuval, a bay in the Bento Gomes River (Mato Grosso), and in a bay in the Acurizal Reserve (Mato Grosso do Sul). Baía Piuval and Acurizal plots contained a total of 22 and 18 macrophyte species, respectively. At both sites Eichhornia azurea and Salvinia auriculata occurred most frequently as dominant or co-dominant species. Chi² analysis, used to quantify zonations along depth gradients, generated four different groups of species ( p < 0.05) for Baía Piuval. For Acurizal, two significantly different groups ( p < 0.05) occurred with an intermediate assemblage of species that could be assigned to either group. Canonical correspondence analysis, used to analyze species distributions, showed a pattern consistent with the Chi² results for Baía Piuval but not for Acurizal. Higher species richness and diversity occurred where dry season and low water levels coincided and richness was generally highest in proximal plots where water depths were lowest. Our results are consistent with the few other plant ecological studies reported for the Pantanal. This study can be considered additive to needed baseline data on biota and ecology of this region of South America.This revised version was published online in March 2005 with corrections to the issue cover date.     

Analyzing the Spatial Structure of Broughtonia cubensis (Orchidaceae) Populations in the Dry Forests of Guanahacabibes,Cuba

A current key issue in ecology is the role of spatial effects on population and community dynamics. In this paper, we test several hypotheses related to spatial structures and coexistence of epiphytic tropical orchid species with special emphasis on the endemic species Broughtonia cubensis. More specifically, we explored the spatial structure of orchid–host plant communities at three different levels of organization (occupied vs. nonoccupied host trees, trees with B. cubensis vs. other orchids, and reproductive vs. nonreproductive B. cubensis plants). We mapped all potential host trees and orchids at three 20 × 20 m plots and applied techniques of spatial point pattern analysis such as mark connection and mark correlation functions to evaluate departures from randomized communities. We found spatial aggregation of trees with epiphytic orchids and segregation between trees with and without epiphytic orchids, and that there was an intraspecific spatial aggregation of B. cubensis in relation to the other seven epiphytic orchid species. Furthermore, we found spatial aggregation of reproductive B. cubensis individuals and segregation between reproductive and nonreproductive individuals on their phorophytes. Thus, orchid–host plant communities show hierarchical spatial structuring with aggregation and segregation at different levels of organization. Our results point to an enhancement of local species in the coexistence of tropical epiphytic orchid communities, by reducing competition through niche differentiation.     

Spatial Pattern of Pindó Palm (Syagrus romanzoffiana) Recruitment in Argentinian Atlantic Forest: The Importance of Tapir and Effects of Defaunation

Hunting pressure, fragmentation and deforestation have caused global declines in animal abundance, and the consequences for plant communities are poorly understood. Many large‐seeded plants, for instance, depend on large and endangered vertebrates for seed dispersal. In some Semi‐deciduous Atlantic Forests, endangered tapirs (Tapirus terrestris) are major dispersers of pindó palms (Syagrus romanzoffiana). Here, we compare recruitment patterns of pindó palms between protected and disturbed (defaunated) Atlantic Forest areas in Argentina and evaluate the potential consequences of the lack of the main disperser for pindó palm regeneration. We analyzed the number and spatial pattern of pindó adults, offspring, and tapir dung piles within ten plots established in an area spanning tapir latrines inside Iguazú National Park and in a fragmented forest area outside the park where tapir is locally extinct. In both areas, we evaluated recruitment levels beneath 24 adult palms in circular plots centered on adult stems. We found lower pindó palm recruitment outside the park where offspring tended to be aggregated around adult palms. In contrast, in Iguazú National Park offspring were spatially associated with tapir dung‐piles, in which most offspring were registered. Recruitment under adults was higher outside the park suggesting a lower rate of seed removal in disturbed areas. Our results show that tapir dispersal promotes higher recruitment levels of pindó offspring and shapes their spatial pattern, breaking the spatial association with adult (presumably maternal) palms. These results are useful for predicting the impact of local tapir extinction on this palm.     

An evaluation of the state of spatial point pattern analysis in ecology

Over the last two decades spatial point pattern analysis (SPPA) has become increasingly popular in ecological research. To direct future work in this area we review studies using SPPA techniques in ecology and related disciplines. We first summarize the key elements of SPPA in ecology (i.e. data types, summary statistics and their estimation, null models, comparison of data and models, and consideration of heterogeneity); second, we review how ecologists have used these key elements; and finally, we identify practical difficulties that are still commonly encountered and point to new methods that allow current key questions in ecology to be effectively addressed. Our review of 308 articles published over the period 1992–2012 reveals that a standard canon of SPPA techniques in ecology has been largely identified and that most of the earlier technical issues that occupied ecologists, such as edge correction, have been solved. However, the majority of studies underused the methodological potential offered by modern SPPA. More advanced techniques of SPPA offer the potential to address a variety of highly relevant ecological questions. For example, inhomogeneous summary statistics can quantify the impact of heterogeneous environments, mark correlation functions can include trait and phylogenetic information in the analysis of multivariate spatial patterns, and more refined point process models can be used to realistically characterize the structure of a wide range of patterns. Additionally, recent advances in fitting spatially‐explicit simulation models of community dynamics to point pattern summary statistics hold the promise for solving the longstanding problem of linking pattern to process. All these newer developments allow ecologists to keep up with the increasing availability of spatial data sets provided by newer technologies, which allow point patterns and environmental variables to be mapped over large spatial extents at increasingly higher image resolutions.     

Effects of harvester ants on plant species distribution and abundance in a serpentine grassland

Seed harvesting ants can have important effects on the composition and structure of plant communities. We investigated two effects of Messor andrei, the black seed-harvesting ant, on a serpentine grassland plant community in northern California. First, to determine if selective seed predation by ants affects plant community composition, we excluded harvester ants from 1-mediameter circular plots of grassland. Abundances of all species on these plots and on control plots were measured before and after exclosure. Second, to determine if M. andrei nest mounds affect plant community composition, we compared plant species abundances on and off nest mounds. M. andrei deposit large amounts of organic matter on their nest mounds over a foraging season, so mounds may alter the edaphic environment. The exclusion of seed-harvesting activity did not cause changes in the plant community. Nest mounds had a strong effect on plant communities: there were many more grasses and fewer forbs on ant mounds, although at least one forb, Lepidium nitidum, produced twice as many seeds when it grew on nest mounds. We found that nest mounds formed islands of higher-temperature soil in the serpentine grassland. Received: 31 March 1997 / Accepted: 6 May 1997     

The decomposition of Shannon's entropy and a confidence interval for beta diversity

Beta diversity is among the most employed theoretical concepts in ecology and biodiversity conservation. Up to date, a self‐contained definition of it, with no reference to alpha and gamma diversity, has never been proposed. Using Kullback‐Leibler divergence, we present the explicit formula of Shannon's β entropy, a bias correction for its estimator and a confidence interval. We also provide the mathematical framework to decompose Shannon diversity into several hierarchical nested levels. From botanical inventories of tropical forest plots in French Guiana, we estimate Shannon diversity at the plot, forest and regional level. We believe this is a complete and usefulness toolbox for ecologists interested in partitioning biodiversity.     

Spatial scale and sampling size affect the concordance between remotely sensed information and plant community discrimination in restoration monitoring

Remote sensing cartography and GIS are part of ordinary practice in restoration ecology in discriminating patches of habitats, defining objectives, and planning the monitoring phase, but derived information is not always consistent with field survey. We assessed the mapping process efficiency in discriminating different communities, relying on plant composition data, and considering the effect of sample size and plot dimension (grain), in a heterogeneous environment in Tuscany (central Italy). We identified four land cover classes on a land cover map produced with object-oriented technique; hence we conducted a sampling of 64 plots (4 zones × 4 classes × 4 plots), estimating vascular plant cover using a point-quadrant method. Plots were nested squares with side lengths of 0.50 m, 1 m and, limited to a sub-sample, 2 m. We evaluated the effect of sample size and grain using permutational multivariate analysis of variance (PERMANOVA), testing the simultaneous response of species composition compared to land cover classes. Results demonstrated that for a sample size of 64 plots, grain does not influence the ability of discriminating among the habitat types investigated, while for a smaller sub-sample the effect of grain is significant and communities cannot be distinguished at all plot dimensions. Outcomes corroborate the hypothesis that sampling at a series of scales of observations and an adequate sample size can improve monitoring efficiency in restoration ecology. Nomenclature: Pignatti (1982) and Conti et al (2005) for Festuca.     

Functional dependence underlies a positive plant-grasshopper richness relationship

A central focus of ecology is identifying the factors that shape spatial patterns of species diversity and this is particularly relevant in an era of global change. Positive relationships between plant and consumer diversity are common, but could be driven by direct responses of each trophic level to underlying environmental gradients, or indirectly where changes in environmental conditions propagate through food webs. Here we use structural equation modeling to examine the relative importance of soil resource availability and disturbance (fire) in mediating relationships between plant and grasshopper richness in insular grasslands. We found a positive relationship between plant and consumer richness that became stronger after accounting for disturbance, despite unique responses of plants and consumers to the two environmental gradients. Plant richness responded to an underlying gradient in soil resource availability. Time since the last fire had a direct positive effect on grasshopper richness but had no effect on plant richness. This work supports that plant and consumer richness are functionally linked, rather than having similar responses to environmental gradients. By disentangling the direct and indirect processes underlying a positive relationship between plant and consumer diversity in a natural system that spans multiple environmental gradients, we demonstrate the importance of investigating biodiversity through explicit multivariate models.     

Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains

Aim  This research aims to understand the factors that shape elevational diversity gradients and how those factors vary with spatial grain. Specifically, we test the predictions of the species–productivity hypothesis, species–temperature hypothesis, the metabolic theory of ecology and the mid-domain effects null model. We also examine how the effects of productivity and temperature on richness depend on spatial grain.
Location  Deciduous forests along an elevational gradient in Great Smoky Mountains National Park, USA.
Methods  We sampled 22 leaf litter ant assemblages at three spatial grains, from 1-m² quadrats to 50 × 50 m plots using Winkler samplers.
Results  Across spatial grains, warmer sites had more species than did cooler sites, and primary productivity did not predict ant species richness. We found some support for the predictions of the metabolic theory of ecology, but no support for the mid-domain effects null model. Thus, our data are best explained by some version of a species–temperature hypothesis.
Main conclusions  Our results suggest that temperature indirectly affects ant species diversity across spatial grains, perhaps by limiting access to resources. Warmer sites support more species because they support more individuals, thereby reducing the probability of local extinction. Many of our results from this elevational gradient agree with studies at more global scales, suggesting that some mechanisms shaping ant diversity gradients are common across scales.