Colonization rules and spatial distributions in ecology

Because they are intuitive and mathematically straight-forward, colonization rules are often used to model spatial patterns in ecology. Colonization rules assign individuals to categories according to the locations of previous colonists. In this note, a compact introduction to colonization rules in ecology is presented with implications for autocorrelation and spatial distributions. I use the colonization rule approach to unify a diverse set of spatial and species diversity analyses, exploring future extensions to incorporate greater realism.     

Explaining variation in tropical plant community composition: influence of environmental and spatial data quality

The degree to which variation in plant community composition (beta-diversity) is predictable from environmental variation, relative to other spatial processes, is of considerable current interest. We addressed this question in Costa Rican rain forest pteridophytes (1,045 plots, 127 species). We also tested the effect of data quality on the results, which has largely been overlooked in earlier studies. To do so, we compared two alternative spatial models [polynomial vs. principal coordinates of neighbour matrices (PCNM)] and ten alternative environmental models (all available environmental variables vs. four subsets, and including their polynomials vs. not). Of the environmental data types, soil chemistry contributed most to explaining pteridophyte community variation, followed in decreasing order of contribution by topography, soil type and forest structure. Environmentally explained variation increased moderately when polynomials of the environmental variables were included. Spatially explained variation increased substantially when the multi-scale PCNM spatial model was used instead of the traditional, broad-scale polynomial spatial model. The best model combination (PCNM spatial model and full environmental model including polynomials) explained 32% of pteridophyte community variation, after correcting for the number of sampling sites and explanatory variables. Overall evidence for environmental control of beta-diversity was strong, and the main floristic gradients detected were correlated with environmental variation at all scales encompassed by the study (c. 100–2,000 m). Depending on model choice, however, total explained variation differed more than fourfold, and the apparent relative importance of space and environment could be reversed. Therefore, we advocate a broader recognition of the impacts that data quality has on analysis results. A general understanding of the relative contributions of spatial and environmental processes to species distributions and beta-diversity requires that methodological artefacts are separated from real ecological differences.     

Small-scale environmental heterogeneity and the analysis of species distributions along gradients

Abstract. The observed distribution of a species along an environmental gradient is strongly affected by environmental variability within a quadrat. Because a quadrat does not represent a point along an environmental gradient, but rather a range of conditions, it is likely to contain species not typically associated with the mean conditions in the quadrat. Systematic relationships exist between a species' true distribution, the observed distribution as a function of mean quadrat environment, and the frequency distribution of the environment within that quadrat. The observed species habitat breadth increases and the observed maximum abundance decreases as within-quadrat environmental heterogeneity increases. If species distributions or beta diversities are to be compared among species or coenoclines, they should be correctedforintra-quadratheterogeneity.Wederive simple corrections for environmental heterogeneity. The distributions of hardwood forest understory species along a soil acidity gradient in the North Carolina piedmont are presented as an example.     

Fine-scale spatial variation in plant species richness and its relationship to environmental conditions in coastal marshlands

Previous studies have shown that variations in environmental conditions play a major role in explaining variations in plant species richness at community and landscape scales. In this study, we considered the degree to which fine-scale spatial variations in richness could be related to fine-scale variations in abiotic and biotic factors. To examine spatial variation in richness, grids of 1 m² plots were laid out at five sites within a coastal riverine wetland landscape. At each site, a 5 × 7 array of plots was established adjacent to the river’s edge with plots one meter apart. In addition to the estimation of species richness, environmental measurements included sediment salinity, plot microelevation, percent of plot recently disturbed, and estimated community biomass. Our analysis strategy was to combine the use of structural equation modeling (path modeling) with an assessment of spatial association. Mantel’s tests revealed significant spatial autocorrelation in species richness at four of the five sites sampled, indicating that richness in a plot correlated with the richness of nearby plots. We subsequently considered the degree to which spatial autocorrelations in richness could be explained by spatial autocorrelations in environmental conditions. Once data were corrected for environmental correlations, spatial autocorrelation in residual species richness could not be detected at any site. Based on these results, we conclude that in this coastal wetland, there appears to be a fine-scale mapping of diversity to microgradients in environmental conditions.     

Community structure in north temperate ants: temporal and spatial variation

Summary Ant communities in Vermont and New York woods were sampled in four time periods to determine species composition, relative abundances, and nest locations in space. The Vermont community was richer, containing more species and higher nest densities than New York. Both communities followed the geometric distribution of species abundances, suggesting that a single resource was mediating competition. The resource most clearly implicated was suitable nest sites, principally pre-formed plant cavities. Nonrandom species associations, underdispersion in every season, and the occurrence of incipient nests overwintering aboveground all implicated shortage of such cavities. Furthermore, microhabitat differences which produce suitable nest sites occur over a very small scale in these communities.     

武钢厂区景观结构与绿地空间布局研究

应用景观生态学原理对武钢厂区景观结构及结地景观的空间布局进行了调查分析,结果表明,构成武钢厂区景观结构的主体是以建筑铺装玻璃块,工业预留地斑块,公路和铁路廊道组成的人工景观要求,其面积占景观总面积的74．37％,班块优势率达57．70％,其中,建筑铺装斑块面积比率最高（56．43％）,优势度最大（35．59％）,专用绿地斑块破碎化指数高（5．0591ind.hm^-2),分布范围广,道路廊道密度大（13．7772km.km^-2),且公路比铁路对景观的影响更大,既体现了工业区景观结构的基本特征,也反映了武钢厂区趋于合理的景观格局,在绿地景观中,道路绿带斑块数目多,面积比率高（66．07％）,优势度最大（75．31％）,以乔木树种为主体的防护林斑块平均面积大（8454．45m2),环境效益高,观赏绿地斑块植物种类丰富,景观效果好,以道路绿带为骨架,成片防护林和观赏绿地为中心,将各分厂绿地接成四大绿化区域的武钢厂区绿地系统已初步形成,但厂区荒草地斑块优势度偏高,防护林面积较小,道路绿带仍不完整,应进一步重视厂区废弃地开发和绿地系统建设。     

Applications of biotope mapping for spatial environmental planning and policy: case studies in urban ecosystems in Korea

The purpose of this paper is to investigate the current status and method of biotope mapping for practical use for landscape planning and environmental policy in urban ecosystem in Korea. We examine current ecological mapping of Seoul, Seongnam, Daegu, and Yongin. Ecological mapping is examined closely in terms of investigation methodology, investigation subject, classification of urban landscape, and the present condition of application. Biotope mapping in Seoul and Seongnam were carried out by the city governments concerned with the pre-set budgets earmarked for mapping. In order to promote the utilization of biotope maps for city planning in Korea, the following actions should be considered. First, the survey method should be standardized by introducing a uniform standard with respect to the scope of survey, the quality of primary data used, the survey method, and the level of the survey. Second, it is necessary to identify a basic category of biotope for each area by consolidating the outcome of the previous surveys. Third, it is highly desirable to minimize the differences between the evaluation criteria and the assessment factors. Fourth, it is ideal to apply the results of the biotope evaluation to city planning in an indirect manner through reflecting the results first in the landscape plans. In order to facilitate this alternative utilization, it is necessary to strengthen the control provisions contained in the ordinances of the city concerned or to enact a set of new provisions in the ordinances so that biotope mapping could be used more widely as a criterion for the spatial environmental impact assessment.     

Distribution of epipelic diatoms in artificial fishponds along environmental and spatial gradients

Although epipelic diatoms play a key role in primary production of many ecosystems, many aspects of their biodiversity, ecology and geographical distribution are poorly understood. The present study is based on sampling of 45 man-made fishponds in the Czech Republic covering an environmental gradient from oligo/dystrophic highland ponds within protected areas to the eutrophic/hypertrophic lowland ponds used for intensive fish production. Diatom distribution patterns assessed using biomass and species composition variables were assessed along environmental and geographical gradients. In total, 185 epipelic diatom taxa were found in the investigated samples. The differences in species composition between sites were correlated with environmental parameters, but not with the geographic distance of the localities. This pattern might suggest that niche-based control, rather than the effect of dispersal limitation, is the main driving force in the species composition of epipelic diatoms in fishponds. The alpha-diversity of sites correlated with altitude, nitrogen and chlorophyll a concentrations but did not correlate with the area of the ponds. The significant relationships between local abundance of species and their regional occupancy were very similar to previous studies of diatoms in boreal streams. In addition, these data concur with patterns known for multicellular organisms suggesting that in this respect diatoms may not differ from groups of organisms with larger body sizes. Handling editor: J. Padisak     

Causes of spatial patterns of dead trees in forest fragments in Illinois

Natural disturbances introduce spatial heterogeneity into forests by causing non-random mortality of trees. We examined whether wind was the primary cause of spatial patterns of dead trees at fragment- and individual tree-levels in three fragments of temperate deciduous forests in Illinois, USA. Dead trees and wind-caused types of mortality were expected to be higher at forest edges, on windward aspects, in poorly-drained soils, and adjacent to existing canopy gaps. The extent of wind-related mortality was determined by comparing spatial and temporal patterns of dead trees, as well as characteristics of trees downed by single windstorms versus all dead trees. At the fragment-level, we used randomly located quadrats of 25×25 m to sample edge and interior areas of Trelease Woods, Brownfield Woods, and Hart Woods in 1995-1996 and again in 1999-2000. We noted type of mortality (standing dead, snapped-off, or uprooted trees), and measured DBH ( 10 cm) and direction of fall of each dead tree. The same measures were made for trees felled by two single storms in 1994. At the individual tree-level, domino effects were evaluated by comparing openness surrounding target treefalls vs. an equal sample size of living trees. The study provided limited evidence that wind caused spatial patterns of dead trees. Instead, spatial patterns of dead trees in the fragments accumulated from domino effects at the individual tree-level in two of the three fragments. Dead trees were more associated with preexisting gaps. Contrary to our predictions at the fragment-level, the frequency of dead trees was not greater at edges, on windward aspects, or in poorly drained soils. This study demonstrated the complexity of spatial patterns of dead trees in forest fragments. The significant domino effects indicated that the occurrence of dead trees was not random, but determined by previous disturbances.     

Simulations to Determine the Mean and Variance of the Point-Object Analogue of the Clark-Evans Statistic

Doguwa and Upton (1988) propose edge corrections for the mean and variance of the average distance X , between a sampling point and the nearest object in a spatial point pattern for the case of a regular grid of sampling points. However, their edge corrections are not applicable to circular sampling windows. This paper extends the range of application of X , by providing its revised mean and variance for use with circular regions.     

Fire acting as an increasing spatial autocorrelation force: Implications for pattern formation and ecological facilitation

Fire is an indissoluble component of ecosystems, however quantifying the effects of fire on vegetation is a challenging task as fire lies outside the typical experimental design attributes. A recent simulation study showed that under increased fire regimes positive tree–tree interactions were recorded (Bacelar et al., 2014). Data from experimental burning plots in an African savanna, the Kruger National Park, were collected across unburnt and annual burn plots. Indices of aggregation and spatial autocorrelation of the distribution of trees between different fire regimes were explored. Results show that the distribution of trees under fire were more clumped and exhibited higher spatial autocorrelation than in unburnt plots. In burnt plots spatial autocorrelation values were positive at finer scales and negative at coarser scales potentially indicating co-existence of facilitation and competition within the same ecosystem depending on the scale. The pattern derived here provides inference for (a) fire acting as an increasing aggregation & spatial autocorrelation force, (b) tree survival under fire regimes is potentially facilitated by forming patches of trees and (c) scale-dependent facilitation and competition coexisting within the same ecosystem with finer scale facilitation and coarser scale competition.     

Edge and shape effects on ant (Hymenoptera: Formicidae) species richness and composition in forest fragments

In this paper, we tested four hypotheses relative to edge and shape effects on ant communities: (i) forest edges have lower species richness than the remnant core; (ii) species richness increases with distance from the edge; (iii) irregularly shaped remnants have lower species richness than more regular remnants; (iv) there is a higher similarity of species composition between edge and core in irregular than in regular remnants. We sampled litter ant communities on the edge and core of ten remnants, in Viçosa, Minas Gerais, Brazil. Species richness was larger at the forest core than at the edges, although did not increase with distance from the edge. Species richness did not vary with shape complexity. The similarity of species composition between edge and core showed a decreasing trend with remnant area, and did not vary with shape complexity. The observed differences of species richness between forest core and edge may be due to higher harshness of edges, caused by environmental changes. The absence of relationship between species richness and distance from the edge might indicate the range of edge effects, which would be smaller than the smallest distance of core sampled. Therefore, edges would affect litter-dwelling ant species richness in a distance smaller than 50 m. The observation of species composition allowed us to notice an effect of fragmentation that would not be noticed if we were considering only species richness. Edge may serve as step to generalist species, which may use it to colonise forest remnants. Furthermore, small remnants are more colonisation-prone by such species, and have a more homogeneous species composition than large remnants.     

A conceptual model of coastal dune ecology synthesizing spatial gradients of vegetation, soil, and geomorphology

Patterns of coastal dune vegetation are closely related with soil conditions controlled by geomorphic forms and processes. This study developed a conceptual model integrating these relationships in a spatially explicit manner. A rectangle of 180 × 280 m containing 126 grids of 20 × 20 m was established in the Sindu coastal dunefield in west Korean Peninsula. Sampling from each grid determined 11 soil properties and identified percent cover of 21 woody and herbaceous plant species. Digital elevation models were generated by topographic survey and used to derive eight topographic parameters. Redundancy analysis and canonical correspondence analysis examined the effect of geomorphic factors on edaphic characteristics and the edaphic influence on spatial distribution of vegetation, respectively. The spatial pattern of soil properties and plant species were inferred from spatial interpolation techniques. In the foredune area, distance from the coastline was a significant indicator of soil nutrients derived from the marine sources by aeolian processes. This favored the dominance by Elymus mollis. Moisture-tolerant species (e.g., Calamagrostis epigeios) had high cover in the acidic soils of dune slacks, which covaried with wetness index, an indirect measure of the depth to the freshwater table. Vegetation–soil interactions (e.g., nitrogen fixation by legumes) were important in secondary dune areas, with topographic effects less significant. Vegetation, soil, and geomorphic factors are closely connected in a causal chain across a whole dune area. Our model thus addresses the importance of integrating foredune, dune slack, and secondary dune into one continuous system.     

Iterative kriging for removing spatial autocorrelation in analysis of forest genetic trials

Conventional analysis of spatially correlated data in inadequately blocked field genetic trials may give erroneous results that would seriously affect breeding decisions. Forest genetic trials are commonly very large and strongly heterogeneous, so adjustments for micro-environmental heterogeneity become indispensable. This study explores the use of geostatistics to account for the spatial autocorrelation in four Pinus pinaster Ait. progeny trials established on hilly and irregular terrains with a randomized complete block design and large blocks. Data of five different traits assessed at age 8 were adjusted using an iterative method based on semivariograms and kriging, and the effects on estimates of variance components, heritability, and family effects were evaluated in relation to conventional analysis. Almost all studied traits showed nonrandom spatial structures. Therefore, after the adjustments for spatial autocorrelation, the block and family × block variance components, which were extremely high in the conventional analysis, almost disappeared. The reduction of the interaction variance was recovered by the family variance component, resulting in higher heritability estimates. The removal of the spatial autocorrelation also affected the estimation of family effects, resulting in important changes in family ranks after the spatial adjustments. Comparison among families was also greatly improved due to higher accuracy of the family effect estimations. The analysis improvement was larger for growth traits, which showed the strongest spatial heterogeneity, but was also evident for other traits such as straightness or number of whorls. The present paper demonstrates how spatial autocorrelation can drastically affect the analysis of forest genetic trials with large blocks. The iterative kriging procedure presented in this paper is a promising tool to account for this spatial heterogeneity.     

Current problems of environmental gradients and species response curves in relation to continuum theory

Abstract. Comparisons of the positions of species on Grimes'C-S-R triangular ordination model with their responses to individual environmental gradients indicates that the C-S-R model does not necessarily predict species ecological behaviour. The importance of the stress, productivity and disturbance gradients relative to other environmental gradients needs to be determined. In studies of species behaviour along a biomass/productivity gradient the collective vegetation property, biomass, has been confused with the environmental factor, fertility. Patterns of responses to biomass gradients e.g. Keddy's centrifugal model, should be examined in a two-dimensional environmental space to avoid such confounding effects. Assumptions regarding the shapes of species responses to environmental gradients remain untested. A recent model of species response functions to environmental gradients suggested that skewed responses curves show a pattern in the direction of the skew, always with the tail towards the presumed most mesic position on the gradient. Further evidence is presented to support this model for a temperature gradient in eucalypt forest in south-eastern Australia. 21 out of 24 species tested conform to the model.     

Experimental biogeography: the role of environmental gradients in high geographic diversity in Cape Proteaceae

One of the fundamental dimensions of biodiversity is the rate of species turnover across geographic distance. The Cape Floristic Region of South Africa has exceptionally high geographic species turnover, much of which is associated with groups of closely related species with mostly or completely non-overlapping distributions. A basic unresolved question about biodiversity in this global hotspot is the relative importance of ecological gradients in generating and maintaining high geographic turnover in the region. We used reciprocal transplant experiments to test the extent to which abiotic environmental factors may limit the distributions of a group of closely related species in the genus Protea (Proteaceae), and thus elevate species turnover in this diverse, iconic family. We tested whether these species have a “home site advantage” in demographic rates (germination, growth, mortality), and also parameterized stage-structured demographic models for the species. Two of the three native species were predicted to have a demographic advantage at their home sites. The models also predicted, however, that species could maintain positive population growth rates at sites beyond their current distribution limits. Thus the experiment suggests that abiotic limitation under current environmental conditions does not fully explain the observed distribution limits or resulting biogeographic pattern. One potentially important mechanism is dispersal limitation, which is consistent with estimates based on genetic data and mechanistic dispersal models, though other mechanisms including competition may also play a role. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Spatial pattern of diversity in an old-growth temperate forest in Northeastern China

Species diversity has attracted particular attention because of its significance for helping determine present species performance and likely future community composition. The spatial pattern of species diversity (species richness, abundance and Shannon diversity) in Changbai temperate forest in Northeastern China was studied to investigate the present and likely causes for the formation of spatial patterns. To fulfill this goal, three aspects of diversity were addressed: 1) changes in the relationships of the diversity variables, species richness, abundance and Shannon diversity, to sampling area and sampling design. The three diversity variables were found to respond to sampling area in a dissimilar way. Sampling design had no significant effect on the diversity variable-area curves. The power function, which was derived under the assumption that the forest was in equilibrium, did not fit the observed species-area curves, indicating that the Changbai temperate forest was probably not in equilibrium. 2) Variograms, used to examine the spatial structure of species diversity, showed that the spatial structure of species diversity in the Changbai temperate forest was weakly anisotropic. 3) Partitioning the variation of species diversity into spatial and environmental factors indicated that the spatial pattern of the Changbai forest community was unpredictable, probably because there were many undetermined processes controlling its development.     

Interacting effects of landscape context and habitat quality on flower visiting insects in agricultural landscapes

Landscape context and habitat quality may have pronounced effects on the diversity of flower visiting insects. We investigated whether the effects of landscape context and habitat quality on flower visiting insects interact in agricultural landscapes in the Netherlands. Landscape context was expressed as the area of semi-natural habitats or the density of linear landscape features, and was quantified at spatial scales ranging from 250 to 2000 m. Habitat quality was determined as flower abundance. Species richness and abundance of hoverflies and bees were determined along 16 stream banks experiencing similar environmental conditions but situated in areas with contrasting landscape context. Only flower abundance and the area of semi-natural habitats within 500–1000 m were significantly related to species richness of hoverflies and bees and these factors had interacting effects on both species groups. Our results suggest that the regional area of semi-natural habitats had a positive effect on hoverfly species richness when flower abundance was relatively high, but not when flower abundance was low. Moreover, flower abundance had positive effects on hoverfly species richness only in areas with relatively many semi-natural habitats. Contrastingly, flower abundance had a more positive effect on bee species richness in landscapes with few semi-natural habitats compared to landscapes with more semi-natural habitats. Our results suggest that the importance of landscape context for the species richness of flower visiting insects depends upon the quality of the habitat patches.     

Variation in spatial and temporal gradients in zooplankton spring development: the effect of climatic factors

1. We examined the temporal and spatial heterogeneity of zooplankton in lake surface waters during the spring of 3 years in Lake Washington, U.S.A., a large lake with a high production of sockeye salmon fry. 2. We show large within‐season and among‐year variation in the horizontal distribution of temperature, chlorophyll a concentration, and zooplankton in the lake. The main pattern, a delay in zooplankton population increase from the north‐ to the south‐end of the lake, recurred in each year and was persistent within each spring. 3. The delay is primarily caused by the development of a temperature gradient during spring warming, as cold mountain water enters the south end of the lake, while warm water enters the north end via a river draining a nearby lake. Climate factors, such as air temperature and precipitation during winter and spring, appear to influence the extent of the delay of zooplankton increase. 4. If the climate continues to warm, the temporal disconnection in zooplankton development between lake areas immediately influenced by cold river inflow and areas that are influenced by spring warming may increase in magnitude. Thus, the different areas of the lake may not contribute equally to fish production.