Abundance and dominance become less predictable as species richness decreases

Aim To test the hypothesis that communities with higher diversity have more predictable properties by examining patterns of community structure along a species richness gradient. Location Trinidad and Tobago (11°00 N, 61°00 W), on the South American continental shelf, opposite the Orinoco River delta, north‐east Venezuela. Methods We used quantile regressions to investigate how three total abundance, absolute and relative dominance measures – numerical abundance, biomass and energy use, respectively – change across a species richness gradient. We investigated which allocation rule best mimics community assembly in this species richness gradient by examining the abundance of the dominant species and comparing it with predictions of niche apportionment models. Results All measures of total abundance increase on average across the gradient, but the upper limit remains constant. On average, absolute dominance is constant, but the distance between the upper and lower limits decreases along the gradient. Relative dominance decreases with species richness. Observed dominance patterns are best described by Tokeshi's random fraction model. Main conclusions Our results show that both total abundance and absolute dominance become increasingly variable as biodiversity decreases. Consequently, our study suggests that ecosystem properties are less predictable when biodiversity is lower.     

Confronting different models of community structure to species-abundance data: a Bayesian model comparison

Species abundances are undoubtedly the most widely available macroecological data, but can we use them to distinguish among several models of community structure? Here we present a Bayesian analysis of species‐abundance data that yields a full joint probability distribution of each model's parameters plus a relatively parameter‐independent criterion, the posterior Bayes factor, to compare these models. We illustrate our approach by comparing three classical distributions: the zero‐sum multinomial (ZSM) distribution, based on Hubbell's neutral model, the multivariate Poisson lognormal distribution (MPLN), based on niche arguments, and the discrete broken stick (DBS) distribution, based on MacArthur's broken stick model. We give explicit formulas for the probability of observing a particular species‐abundance data set in each model, and argue that conditioning on both sample size and species count is needed to allow comparisons between the two distributions. We apply our approach to two neotropical communities (trees, fish). We find that DBS is largely inferior to ZSM and MPLN for both communities. The tree data do not allow discrimination between ZSM and MPLN, but for the fish data ZSM (neutral model) overwhelmingly outperforms MPLN (niche model), suggesting that dispersal plays a previously underestimated role in structuring tropical freshwater fish communities. We advocate this approach for identifying the relative importance of dispersal and niche‐partitioning in determining diversity of different ecological groups of species under different environmental conditions.     

Ecological niche models display nonlinear relationships with abundance and demographic performance across the latitudinal distribution of Astragalus utahensis (Fabaceae)

The potential for ecological niche models (ENMs) to accurately predict species' abundance and demographic performance throughout their geographic distributions remains a topic of substantial debate in ecology and biogeography. Few studies simultaneously examine the relationship between ENM predictions of environmental suitability and both a species' abundance and its demographic performance, particularly across its entire geographic distribution. Yet, studies of this type are essential for understanding the extent to which ENMs are a viable tool for identifying areas that may promote high abundance or performance of a species or how species might respond to future climate conditions. In this study, we used an ensemble ecological niche model to predict climatic suitability for the perennial forb Astragalus utahensis across its geographic distribution. We then examined relationships between projected climatic suitability and field‐based measures of abundance, demographic performance, and forecasted stochastic population growth (λ_s). Predicted climatic suitability showed a J‐shaped relationship with A. utahensis abundance, where low‐abundance populations were associated with low‐to‐intermediate suitability scores and abundance increased sharply in areas of high predicted climatic suitability. A similar relationship existed between climatic suitability and λ_s from the center to the northern edge of the latitudinal distribution. Patterns such as these, where density or demographic performance only increases appreciably beyond some threshold of climatic suitability, support the contention that ENM‐predicted climatic suitability does not necessarily represent a reliable predictor of abundance or performance across large geographic regions.     

Stochastic determinants of assemblage patterns in coral reef fishes: a quantification by means of two models

Synopsis One perspective emphasizing the importance of stochastic processes in determining coral reef fish assemblages implies that there is little organization in species richness, abundance structure, and spatial distribution. We examine the degree to which this perspective is correct by analyzing distribution of fishes on a collection of patch reefs (Discovery Bay, Jamaica). We ask the question whether these patches accumulate species and individuals in a manner consistent with stochastic expectations. To address this question we use two conceptual models, each permitting a different insight. One assumes that fish are distributed stochastically on patches while the other assumes presence of restrictions on fish distribution due to habitat structure. For each conceptual model we use two types of benchmark: we compare observed patterns to those predicted by theoretical models, and we also compare observed patterns to those obtained from a random reallocation of fish individuals to patches. We found that the conceptual model assuming stochastic processes appeared to provide weaker explanation of patterns than the conceptual model that includes restrictions due to habitat structure. Further, and more importantly, we found that (i) the community is shaped by a mixture of stochastic and non-stochastic mechanisms, and (ii) the stochastic assembly processes decrease in importance for species restricted to fewer microhabitat types and sites. Our study therefore indicates that patches do accumulate individuals and species in a manner consistent with stochastic expectations, however, this applies primarily to the habitat generalists (unrestricted species). By the same token, increased habitat specialization by some species imposes constraints on the stochastic model such that it eventually fails.     

Metabolic constraints on the body size scaling of extreme population densities

Pest outbreaks, harmful algal blooms and population collapses are extreme events with critical consequences for ecosystems. Therefore, understanding the ecological mechanisms underlying these extreme events is crucial. We evaluated theoretical predictions on the size scaling and variance of extreme population abundance by combining (i) the generalized extreme value (GEV) theory and (ii) the resource-limited metabolic restriction hypothesis for population abundance. Using the phytoplankton data from the L4 station in the English Channel, we showed a negative size scaling of the expected value of maximal density, whose confidence interval included the predicted metabolic scaling (α = −1) supporting theoretical predictions. The role of resources and temperature in the distribution of the size–abundance pattern and residuals was well characterized by the GEV distribution. This comprehensive modelling framework will allow to elucidate community structure and fluctuations and provide unbiased return times estimates, thereby improving the prediction accuracy of the timing of the population outbreaks.     

Influence of Habitat Structure on Pomacentrus sulfureus, A Western Indian Ocean Reef Fish

The influence of habitat on the distribution and abundance of Pomacentrus sulfureus was investigated on coral reefs in Zanzibar. Fish censuses were conducted using the simultaneous belt-transect method and substrate data were gathered using a point-base method. Densities of juvenile and adult P. sulfureus were examined in relation to habitat composition and structure. The influence of habitat structure on coral reef fishes remains debated and has been obscured by the various methods, scales and levels of detail that have been used. In this study, we compared two measures of structural complexity. One was a contour measure (rugosity) and the other was the percent cover of branching structures. Both were applied on the same scale but differed in the level of detail. P. sulfureus was distributed in an uneven pattern around Zanzibar Island and the distribution appeared to reflect local and regional differences in habitat structure. Multiple regression models identified relations between juvenile and adult P. sulfureus abundance and specific habitat features. The majority of the variation in juvenile numbers was attributed to percent cover of branching structures, the high-resolution measure. However, adult abundance was unrelated to this measure. Rugosity, the low-resolution measure of structural complexity, appeared to influence neither adult nor juvenile P. sulfureus. The best predictor of adult abundance was substrate diversity (inverse relationship), which, however, did not contribute to the proportion of explained variation in juvenile abundance. In addition, there was a weak positive relationship between adult abundance and the number of hard coral growth forms present. The results indicate that P. sulfureus exhibit an ontogenetic shift in habitat use. However, the regression models also revealed that adults per se have a positive effect on juvenile numbers. We suggest that the limited habitat use of juvenile P. sulfureus is contained within that of conspecific adults, and that juveniles are likely to adopt more general habitat associations as they grow.     

Relationships between distribution and abundance vary with spatial scale and ecological group in stream bryophytes

1. We examined whether the local abundance of stream bryophytes in a boreal drainage basin (Koutajoki system in northeastern Finland) correlated with their: (i) regional occupancy; (ii) provincial distribution in northwestern Europe; and (iii) global range size. We specifically tested whether aquatic and semi‐aquatic species differ in their distribution–abundance relationships. We also analysed the frequency distributions of occupancy at two spatial scales: within the focal drainage system and across provinces of northwestern Europe. 2. Regional occupancy and mean local abundance of stream bryophytes were positively correlated, and the relationship was rather strong in aquatic species but very weak in semi‐aquatic species. Local abundance was related neither to provincial distribution nor global distribution. 3. Species frequency distributions differed between regional occupancy and provincial distribution. While most species were rare with regard to their regional occupancy within the focal drainage system, most of the same set of species were common and occurred in most provinces in northwestern Europe. 4. The results indicate the presence of dominants (core species) and transients/subordinates (satellite species) among stream bryophytes, highlighting marked differentiation in life‐history strategies and growth form. The observed abundance–occupancy relationships suggest that dispersal limitation and metapopulation processes may govern the dynamics of obligatory aquatic stream bryophytes. In semi‐aquatic species, however, habitat availability may be more important in contributing to regional occupancy.     

Motif frequency and evolutionary search times in RNA populations

RNA molecules, through their dual identity as sequence and structure, are an appropriate experimental and theoretical model to study the genotype-phenotype map and evolutionary processes taking place in simple replicator populations. In this computational study, we relate properties of the sequence-structure map, in particular the abundance of a given secondary structure in a random pool, with the number of replicative events that an initially random population of sequences needs to find that structure through mutation and selection. For common structures, this search process turns out to be much faster than for rare structures. Furthermore, search and fixation processes are more efficient in a wider range of mutation rates for common structures, thus indicating that evolvability of RNA populations is not simply determined by abundance. We also find significant differences in the search and fixation processes for structures of same abundance, and relate them with the number of base pairs forming the structure. Moreover, the influence of the nucleotide content of the RNA sequences on the search process is studied. Our results advance in the understanding of the distribution and attainability of RNA secondary structures. They hint at the fact that, beyond sequence length and sequence-to-function redundancy, the mutation rate that permits localization and fixation of a given phenotype strongly depends on its relative abundance and global, in general non-uniform, distribution in sequence space.     

Relative importance of the land‐use composition and intensity for the bird community composition in anthropogenic landscapes

Humans are changing the biosphere by exerting pressure on land via different land uses with variable intensities. Quantifying the relative importance of the land‐use composition and intensity for communities may provide valuable insights for understanding community dynamics in human‐dominated landscapes. Here, we evaluate the relative importance of the land‐use composition versus land‐use intensity on the bird community structure in the highly human‐dominated region surrounding Paris, France. The land‐use composition was calculated from a land cover map, whereas the land‐use intensity (reverse intensity) was represented by the primary productivity remaining after human appropriation (NPP_remaining), which was estimated using remote sensing imagery. We used variance partitioning to evaluate the relative importance of the land‐use composition versus intensity for explaining bird community species richness, total abundance, trophic levels, and habitat specialization in urban, farmland, and woodland habitats. The land‐use composition and intensity affected specialization and richness more than trophic levels and abundance. The importance of the land‐use intensity was slightly higher than that of the composition for richness, specialization, and trophic levels in farmland and urban areas, while the land‐use composition was a stronger predictor of abundance. The intensity contributed more to the community indices in anthropogenic habitats (farmland and urban areas) than to those in woodlands. Richness, trophic levels, and specialization in woodlands tended to increase with the NPP_remaining value. The heterogeneity of land uses and intensity levels in the landscape consistently promoted species richness but reduced habitat specialization and trophic levels. This study demonstrates the complementarity of NPP_remaining to the land‐use composition for understanding community structure in anthropogenic landscapes. Our results show, for the first time, that the productivity remaining after human appropriation is a determinant driver of animal community patterns, independent of the type of land use.     

Long-term demographic surveys reveal a consistent relationship between average occupancy and abundance within local populations of a butterfly metapopulation

Species distribution models are the tool of choice for large-scale population monitoring, environmental association studies and predictions of range shifts under future environmental conditions. Available data and familiarity of the tools rather than the underlying population dynamics often dictate the choice of specific method – especially for the case of presence–absence data. Yet, for predictive purposes, the relationship between occupancy and abundance embodied in the models should reflect the actual population dynamics of the modelled species. To understand the relationship of occupancy and abundance in a heterogeneous landscape at the scale of local populations, we built a spatio-temporal regression model of populations of the Glanville fritillary butterfly Melitaea cinxia in a Baltic Sea archipelago. Our data comprised nineteen years of habitat surveys and snapshot data of land use in the region. We used variance partitioning to quantify relative contributions of land use, habitat quality and metapopulation covariates. The model revealed a consistent and positive, but noisy relationship between average occupancy and mean abundance in local populations. Patterns of abundance were highly variable across years, with large uncorrelated random variation and strong local population stochasticity. In contrast, the spatio-temporal random effect, habitat quality, population connectivity and patch size explained variation in occupancy, vindicating metapopulation theory as the basis for modelling occupancy patterns in fragmented landscapes. Previous abundance was an important predictor in the occupancy model, which points to a spillover of abundance into occupancy dynamics. While occupancy models can successfully model large-scale population structure and average occupancy, extinction probability estimates for local populations derived from occupancy-only models are overconfident, as extinction risk is dependent on actual, not average, abundance.     

揭示群落结构及其环境响应的联合物种分布模型的研究进展

物种分布模型通常用于基础生态和应用生态研究,用来确定影响生物分布和物种丰富度的因素,量化物种与非生物条件的关系,预测物种对土地利用和气候变化的反应,并确定潜在的保护区.在传统的物种分布模型中,生物的相互作用很少被纳入,而联合物种分布模型(JSDMs)作为近年提出的一种新的可行方法,可以同时考虑环境因素和生物交互作用,因而成为分析生物群落结构和种间相互作用过程的有力工具.JSDMs以物种分布模型(SDMs)为基础,通常采用广义线性回归模型建立物种对环境变量的多变量响应,以随机效应的形式获取物种间的关联,同时结合隐变量模型(LVMs),并基于Laplace近似和马尔科夫蒙脱卡罗模拟的最大似然估计或贝叶斯方法来估算模型参数.本文对JSDMs的产生及理论基础进行归纳总结,重点介绍了不同类型JSDMs的特点及其在现代生态学中的应用,阐述了JSDMs的应用前景、使用过程中存在的问题及发展方向.随着对环境因素与多物种种间关系研究的深入,JSDMs将是今后物种分布模型研究的重点.     

The scale of resource specialization and the distribution and abundance of lycaenid butterflies

Numerous hypotheses have been proposed for the commonly observed, positive relationship between local abundance and geographic distribution in groups of closely related species. Here I consider how hostplant specialization and abundance affect the relative abundance and distribution of lycaenid butterflies (Lepidoptera: Lycaenidae). I first discuss three components of specialization: local specialization, turnover of specialization across a species’ range, and the minimum number of resources (or habitats) required by a species. Within this framework, I then consider one dimension of a lycaenid species’ niche, larval hostplant specialization. In a subalpine region of Colorado, I surveyed 11 lycaenid species and their hostplants at 17 sites. I compare this local information to continental hostplant use and large-scale distributions of the lycaenids and their hostplants. Local abundance of a lycaenid species is positively correlated with its local distribution (the number of sites occupied), but not with its regional or continental distribution. Neither local specialization (the number of hostplants used within one habitat) nor continental specialization (the number of hostplants used across many habitats) is correlated with local lycaenid abundance. Continental specialization is positively correlated with a species’ continental distribution, however. Finally, while generalist butterflies tend to have more hostplant available to them, differences in resource availability do not explain the differences in butterfly abundance. Although local abundance is correlated only with local distribution, I suggest that abundance-distribution relationships might emerge at regional and continental scales if local abundance were averaged across many habitat types. Consideration of the scale of a species’ resource specialization (within or among habitats) appears to be key to understanding the relationships between resource specialization, resource availability, and a species’ abundance and distribution. Received: 1 September 1999 / Accepted: 12 December 1999     

东灵山地区大步甲属物种分布和季节变化的多样性格局

针对东灵山地区的11种不同生境,利用巴氏罐诱法研究大步甲属（Carabus）种类多样怀及其变化与生境的关系,结果表明：（1）本地区有大步甲10种,其中C.manifestus Kraatz和C.crassesculptus Kraatz为优势种,占总数的70.8％,与C.smaragdinus Fischer、C.vladimirskyi Dejean和C.sculptipennis Chaudoir一起构成本地区的常见种;（2）从物种丰富度和数量看,由高到低依次为小龙门林区（以森林为主）、梨园岭退耕区（以灌丛和萌生丛为主）、东灵山主峰区（高海拔,亚高山植物）;物种多样性指数（H‘）由高到低依次为梨园岭退耕区、小龙门林区、东灵山主峰区;均匀度指数值（E）梨园岭退耕区最高,其次为东灵山主峰区,再次为小龙门地区;（3）利用物种的发生与频度数据,对11种不同生境进行栖息地相似性聚类分析,发现东灵山主峰区的3种生境可归为一类,小龙门林区5种生境可归为另一类,梨园岭退耕区的辽东栎萌生丛单独为一类,梨园岭退耕区的山杏和荆条灌丛构成一类,这反映了大步甲属种类分布的地域及生境特点,特别是生境内的湿度、温度和地表覆盖层等特点;（4）通过物种间的相关分析看出：C.smaragdinus和C.granulatus呈显著的正相关关系,反映了它们对退耕区的选择倾向;C.crassesculptus、C.canaliculatus、C.manifestus和Carabus sp.之间,虽然后两种间未达到显著性,但彼此呈正相关关系,反映了这些物种对森林生境选择倾向;（5）大步甲成虫的季节性活动主要在4-9月份间,高峰期集中在6-8份月,各物种在生境内有不同的季节变化趋势,与该地区季节性气候变化一致。研究表明,大步甲物种分布受生境内的湿度、温度、地表覆盖层等环境因素影响,物种多样性与生境肥干扰程度有关,生态恢复将对物种多样性保护具有重要意义。     

吉林蛟河阔叶红松林物种多度分布模型研究

以吉林蛟河阔叶红松林的木本植物为研究对象,将30hm²的样地面积划分为5m×5m,10m×10m,20m×20m,25m×25m的连续取样单元,在4个不同尺度下分别统计各物种在每个取样单元中的有无,得到每个物种在不同尺度下的取样单元数。利用随机分布模型和负二项分布模型分析物种的多度分布,对比预测多度与观测多度讨论两个模型的科学性与实用性。结果表明：对于阔叶红松林而言,负二项分布模型在所有研究尺度上的预测精度都要优于随机分布模型。随机分布和负二项分布的模型预测误差随着研究尺度的增大而增大,因此选取较小的取样单元可以切实提高物种多度的预测精度。利用随机分布和负二项分布模型对多度较小的物种进行预测的效果要优于多度较大的物种。负二项分布模型适合用来模拟阔叶红松林的物种多度分布格局,并且模型的拟合效果受取样单元大小影响。     

Species Abundance in a Forest Community in South China: A Case of Poisson Lognormal Distribution

Case studies on Poisson lognormal distribution of species abundance have been rare, especially in forest communities. We propose a numerical method to fit the Poisson lognormal to the species abundance data at an evergreen mixed forest in the Dinghushan Biosphere Reserve, South China. Plants in the tree, shrub and herb layers in 25 quadrats of 20 m×20 m, 5 m×5 m, and 1 m×1 m were surveyed. Results indicated that: (i) for each layer, the observed species abundance with a similarly small median, mode, and a variance larger than the mean was reverse J-shaped and followed well the zero-truncated Poisson lognormal;(ii) the coefficient of variation, skewness and kurtosis of abundance, and two Poisson lognormal parameters (σ andμ) for shrub layer were closer to those for the herb layer than those for the tree layer; and (iii) from the tree to the shrub to the herb layer, the σ and the coefficient of variation decreased, whereas diversity increased. We suggest that: (i) the species abundance distributions in the three layers reflects the overall community characteristics; (ii) the Poisson lognormal can describe the species abundance distribution in diverse communities with a few abundant species but many rare species; and (iii) 1/σ should be an alternative measure of diversity.     

Quantitative and qualitative beta diversity measures lead to different insights into factors that structure microbial communities

The assessment of microbial diversity and distribution is a major concern in environmental microbiology. There are two general approaches for measuring community diversity: quantitative measures, which use the abundance of each taxon, and qualitative measures, which use only the presence/absence of data. Quantitative measures are ideally suited to revealing community differences that are due to changes in relative taxon abundance (e.g., when a particular set of taxa flourish because a limiting nutrient source becomes abundant). Qualitative measures are most informative when communities differ primarily by what can live in them (e.g., at high temperatures), in part because abundance information can obscure significant patterns of variation in which taxa are present. We illustrate these principles using two 16S rRNA-based surveys of microbial populations and two phylogenetic measures of community beta diversity: unweighted UniFrac, a qualitative measure, and weighted UniFrac, a new quantitative measure, which we have added to the UniFrac website (http://bmf.colorado.edu/unifrac). These studies considered the relative influences of mineral chemistry, temperature, and geography on microbial community composition in acidic thermal springs in Yellowstone National Park and the influences of obesity and kinship on microbial community composition in the mouse gut. We show that applying qualitative and quantitative measures to the same data set can lead to dramatically different conclusions about the main factors that structure microbial diversity and can provide insight into the nature of community differences. We also demonstrate that both weighted and unweighted UniFrac measurements are robust to the methods used to build the underlying phylogeny.     

Small- and broad-scale patterns of distribution of the upper-shore littorinid Nodilittorina pyramidalis in New South Wales

Abstract mall-scale (within-shore) and broad-scale (among-shore) patterns of distribution, abundance and size structure of Nodilittorina pyramidalis were measured on a number of shores in New South Wales. This species showed significant patchiness in distribution and size structure. The mean density and mean shell length of snails differed among patches of shore only a few metres apart, among different heights on the shore and from shore to shore. Differences in densities appeared to be strongly correlated to the distribution of particular microhabitats, especially cracks, crevices and pits within the rock surface. This relationship and the presence or absence of these microhabitats appeared to determine the patterns of distribution of this species within and among different shores. The sizes of snails were not correlated with densities of Nodilittorina nor with the coexisting species, Littorina unifasciata, in the populations examined. Processes influencing the distribution and abundance of Nodilittorina were not investigated, but the patterns of distribution found on these shores suggest that recruitment was limited to areas of topographic complexity, ft also seems likely that limited dispersal by the adult snails retain them within restricted patches on the shore.     

Distribution of abundance across the range in eastern North American trees

Aim  We analysed spatial datasets of abundance across the entirety, or near entirety, of the geographical ranges of 134 tree species to test macroecological hypotheses concerning the distribution of abundance across geographical ranges.
Location  Our abundance estimates came via the USDA Forest Service Forest Inventory and Analysis Eastwide Database, which contains data for 134 eastern North American tree species.
Methods  We extracted measures of range size and the spatial location of abundance relative to position in the range for each species to test four hypotheses: (a) species occur in low abundance throughout most of their geographical range; (b) there is a positive interspecific relationship between abundance and range size; (c) species are more abundant in the centre of their range; and (d) there is a bimodal distribution of spatial autocorrelation in abundance across a species range.
Results  Our results demonstrate that (a) most species (85%) are abundant somewhere in their geographical range; (b) species achieving relatively high abundance tend to have larger range sizes; (c) the widely held assumption that species exhibit an 'abundant-centre distribution' is not well supported for the majority of species; we suggest 'abundant-core' as a more suitable term; and (d) there is no evidence of a bimodal distribution of spatial autocorrelation in abundance.  

Main Conclusions 

For many tree species, high abundance can be achieved at any position in the range, though suitable sites are found with less frequency towards range edges. Competitive relationships may be involved in the distribution of abundance across tree ranges and species with larger ranges (and possibly broader niches) may be affected more by biotic interactions than smaller ranging species.     

The wealth of species: ecological communities, complex systems and the legacy of Frank Preston

General statistical patterns in community ecology have attracted considerable recent debate. Difficulties in discriminating among mathematical models and the ecological mechanisms underlying them are likely related to a phenomenon first described by Frank Preston. He noted that the frequency distribution of abundances among species was uncannily similar to the Boltzmann distribution of kinetic energies among gas molecules and the Pareto distribution of incomes among wage earners. We provide additional examples to show that four different 'distributions of wealth' (species abundance distributions, species–area and species–time relations, and distance decay of compositional similarity) are not unique to ecology, but have analogues in other physical, geological, economic and cultural systems. Because these appear to be general statistical patterns characteristic of many complex dynamical systems they are likely not generated by uniquely ecological mechanistic processes.     

Fungal diversity in ectomycorrhizal communities: sampling effort and species detection

A number of recent review articles on ectomycorrhizal (ECM) fungal community diversity have highlighted the unprecedented increase in the number of publications on this ecologically important but neglected area. The general features of these species-rich, highly dynamic and complex communities have been comprehensively covered but one aspect crucial to our assessment of diversity, namely the sampling of ECM communities has received less attention. This is a complex issue with two principal components, the physical sampling strategy employed and the life cycle traits of the ECM fungi being examined. Combined, these two components provide the image that we perceive as ECM diversity. This contribution will focus primarily on the former of these components using a recent study from a pine forest in central Sweden to highlight some sampling problems and also to discuss some features common to ECM communities. The two commonly used elements of diversity, species richness and community evenness, present rather different problems in the assessment of ECM diversity. The applicability of using current measures of abundance (number or percentage of root tips colonised) to determine community evenness is discussed in relation to our lack of knowledge on the size of individual genets of ECM fungi. The inherent structure of most ECM communities, with a few common species and a large number of rare species, severely limits our ability to accurately assess species richness. A discussion of theoretical detection limits is included that demonstrates the importance of the sampling effort (no. of samples or tips) involved in assessing species richness. Species area abundance plots are also discussed in this context. It is suggested that sampling strategy (bulk samples versus multiple collections of single tips) may have important consequences when sampling from communities where root tip densities differ. Finally, the need for studies of the spatial distribution of ECM on roots in relation to small-scale soil heterogeneity and of the temporal aspects of ECM community dynamics is raised.