Time and space in the community structure of a species-rich limestone grassland

Abstract. The community structure of a species-rich grassland was investigated at a small spatial scale (0.001 m²), to determine whether evidence suggesting assembly rules could be found in temporal or spatial variation in either species richness or guild proportions. The community was alvar limestone grassland on the island of Öland, Sweden. Three sites were sampled: two lightly grazed, the other recently ungrazed. Plots with and without fertilizer were compared. Evidence was sought for restriction on the ability of species to co-occur within a limited spatial area. Restriction due to a limited number of niches available,‘niche limitation’, could be manifest as lower variance in quadrat richness than expected under a null model (i.e. RV_r, the ratio of observed: expected variance in richness, would be < 1.0). In several cases, RV_r values were significantly < 1.0, even using a patch model to allow for possible spatial variation in the environment. Low RV_r values were found only at the smallest square quadrat size, 10 cm². On Fertilized plots in the years immediately after application of fertilizer, low RV_r could not be demonstrated. Explanations of low RV_r other than niche limitation are considered, such as environmental heterogeneity (present and/or historical) and limitations to the co-occurrence of individual plant modules. Assembly rules based on guild membership were sought by looking at the variance across quadrats in the proportions of species from morphological guilds. An assembly rule would be seen as relatively constant proportion, estimated via RV_gp, the ratio of observed: expected variances in guild proportions. Significant guild proportionality was found in some cases. There was no evidence of guild proportionality in the years after the application of fertilizer. The significant effects in RV_r were more numerous than expected on a random basis, though not observed in every site in every year. Similar trends were seen in RV_gp. At the space/time scales examined, the species in a plant community may be constrained by assembly rules only intermittently, e.g. when resources are more limiting (Wiens 1977). Under this concept, when competition is relaxed, such as following fertilizer application, there is a temporary microhabitat ‘waterhole’ in which more species can coexist, and the assembly rules break down, at least temporarily whilst the species composition adjusts. There was some indication of a return to more deterministic community structure four years after fertilization commenced. Variants of van der Maarel's Carousel model were tested. A Niche-limited Carousel Model (i.e. a model in which there is some limitation in the number of species that can occupy a microsite) would imply restricted variation in richness through time for a single quadrat (temporal RV_r). Overall differences between years in species richness were demonstrated, and their effect removed; after this adjustment there was support for the Niche-limited Carousel Model. The extent of this limitation varied between sites. There were also consistent differences between quadrats in species richness. There was little evidence for constancy of guild proportions through time. The site that showed the strongest community structure in time and space, least year-to-year variation in mean species richness, and least response to fertilizer perturbation, is that on the shallowest soil. Possibly the thin soil results in greater resource limitation, supporting suggestions that assembly rules are stronger when resources are more limited.     

Evidence for constraint on species coexistence in vegetation of the Park Grass experiment

Repeated patterns, of a type that would be expected to result from limitations to species coexistence (i.e. assembly rules) were sought in the Park Grass experiment. This classical grassland experiment was sampled in two years, using replicated biomass samples. Variance in a number of measures was examined, and compared to the variance expected under appropriate null models, the latter based on assumptions of no interactions between species. In each case, an assembly rule would result in low variance. Examining variance in species richness between quadrats within a treatment, there was no indication of constraint on species co-occurrences; variance in richness was actually greater than expected under the null model, attributable to environmental variation or perhaps positive interactions between species. However, there was control on biomass, evidenced by variance in total biomass (i.e. over all species) within a treatment being significantly lower than expected under the null model. There was no indication of community structure based on guilds (i.e. functional types). Although there was in 1991 some, non-significant, indication of a constant proportion of species from the legume guild, there was no sign of such an effect in 1992. Searches for intrinsic guilds failed to converge. There was no indication at all of constancy in the proportional representation of guilds by biomass. Thus, there is good evidence for competitive control on plant growth, but none for control of species occurrences. There is no convincing evidence for guild structure in this community at the scale sampled. Possible conflict is discussed between the existence of evidence for temporal stability but the absence of evidence for spatial uniformity. It is concluded that most of the mechanisms proposed for temporal stability will not necessarily lead to control on spatial variation. For many mechanisms, this would depend on the spatial scale examined.     

Evidence for community assembly constraints during succession in dune slack plant communities

Community assembly during succession can be constrained by both local and regional factors. Despite an increasing regional species pool size during succession, we found a limit on the number of species in 1 × 1 m plots in dune slacks. Three alternative hypotheses (habitat heterogeneity, dispersal limitation and niche limitation) explaining this community saturation were tested. A null model analysis showed that species richness in the plots had an unusually low variance suggesting that beta habitat diversity was not likely to explain the limitation on species richness. Because we did not find a correlation between the distribution of species over the slack and their dispersal capacity, we also excluded the dispersal limitation hypothesis. Finally, a guild proportionality analysis revealed that the abundances of forb, graminoid and ruderal species showed low an unusually low variance over all age classes involved. This provides evidence for nonrandom community assembly during succession, likely to be determined by competitive exclusion among species of the same guild.     

Community structure and assembly rules in a dune slack: Variance in richness,guild proportionality,biomass constancy and dominance/diversity relations

There is still disagreement as to whether plant communities can be described as conforming to certain assembly rules. We searched for such rules in the vegetation of four dune slacks in west Wales by examining variance in richness, guild proportionality, biomass constancy and dominance/diversity relations. To test for the occurrence of niche limitation, variance in quadrat species-richness was compared to null models of species occurrence. Using an Overall-model, one site showed a deficit of variance, but this disappeared when a Patch-model analysis was used. It therefore seems likely that the effect observed was due to equality in the species pools adapted to different microenvironments. The proportional representation of functional guilds was examined, in guilds based on morphology and life history. Significant constancy of guild proportions was seen for the annual guild in the only site in which it occurred, perhaps reflecting a regeneration niche. Total biomass per quadrat was examined, and compared to a model in which the biomass of each species was determined at random from the biomasses of that species at the site. Surprisingly, the observed biomasses showed no departure from the null model. However, there was considerable discrepancy from an alternative null model in which biomasses were allocated at random irrespective of species. This would imply that competition limits species coexistence, rather than performance. Examination of the proportions of species from different potential-size classes in each quadrat gave limited confirmation. The local abundance of species (mean biomass) was significantly correlated with the frequency of the species within a site, parallelling the geographical-range/local-abundance correlation previously reported. Five dominance/diversity models were fitted to the biomass data. The best fits were given by the Geometric or the General Lognormal models, though which fitted best varied between and within sites. Between sites, there was some indication of a consistent difference in Preston's gamma. Most of the results are compatible with Gleason's Individualistic concept of the plant community, though there are several pointers that assembly rules have some impact on community structure.     

Guilds and assembly rules in lawn communities

Abstract. The community structure of 11 lawn sites in New Zealand and Fiji was examined in terms of guilds, seeking assembly rules based on guild proportionality. First, associations were analysed, using a new patch model which examined the mean of associations within patches of about 4 cm x 4 cm. As expected from the previously-demonstrated existence of niche limitation at this scale in lawns, the majority of associations between individual species were negative. Even in a lawn only four months old there were significant associations (mostly negative, and one positive). At some sites those species with the most negative associations were those whose morphology might be expected to restrict co-existence with other species. Much ecological theory is based on the assumption that there are limitations to coexistence, related to the resource usage of the species, such that species that are too similar in resource use are less likely to coexist. This theory was tested by looking for evidence of guild proportionality. We defined two guilds, graminoids and forbs. Using these guilds, three of the sites showed significant evenness in proportional representation from the two guilds in quadrats containing four species, i.e. there was less variation in the graminoid:forb ratio than expected on a random basis. Inclusion of species-poor quadrats in a joint analysis over all richness categories overwhelmed this effect, though there was still a strong trend for two of the New Zealand sites: the two species-rich grass lawns. Preliminary analyses of alternative guild delimitations gave no indication that other guild delimitations would have revealed stronger community structure. There was a strong and significant tendency in most sites for the graminoid guild to be more strongly represented in species-poor quadrats, an effect caused by species frequencies.     

Variation in species richness on small grassland quadrats: niche structure or small-scale plant mobility?

Abstract. In this study we make clear that the significance of deviations from an expected variance in species richness as demonstrated in alvar grassland, is a function of spatial pattern at the scale richness was measured, i.e. 10 cm². If corrected for spatial dependence, more than half of the significant cases become nonsignificant. As regards cases of significant variance deficit, which has been interpreted as an indication of niche limitation, we suggest that there may be a simple physical limitation to the number of species on the scale of observation in the form of a low number of plant units which can find a place because of the modal plant size of the species involved. In most cases of significant variance deficit the modal size of the plant units involved was probably bigger than average. Insofar as a significant variance deficit can be demonstrated, the resulting species richness distribution curve should be analyzed and we propose a skewness test, enabling us to differentiate between significantly left-skewed curves (‘niche limitation’), significantly right-skewed curves (‘niche facilitation’) and symmetrical curves. We present results obtained with the G-test, a log-likelihood ratio goodness of fit test. Only few cases of significantly left-skewed curves and a majority of symmetrical curves were found. Attempts to demonstrate guild proportionality in grasslands suffer from the heterogeneity of usually distinguished guilds, such as annuals vs. perennials or graminoids vs. dicotyledons. We observe that niche limitation may occur in alvar grassland, but indications for niche facilitation are stronger. Finally, we conclude that deviations in species richness variance are interesting indications of community structure, but only of spatial structure. Niche structure resulting from assembly rules should be investigated through experiments.     

What constitutes evidence of community structure? A reply to van der Maarel,Noest & Palmer

Abstract. Van der Maarel et al. (1995) — hereafter VNP — criticize our analyses of Öland limestone grassland. They have four general objections, addressed below. 1. VNP attribute our significant results to randomness (‘volatility’); however, significant deviation from randomness cannot be explained by randomness. VNP's conclusion of volatility derives from results that are inconsistent with ours. Their ecological interpretation assumes spatial and temporal near-constancy in the vegetation; we demonstrate that these assumptions are not correct. 2. We discussed physical limitations to plant module packing. VNP give estimated module sizes. We appreciate this information, though data on actual module sizes and overlap patterns would be required before the real module packing effects could be determined. If module packing were really the main cause of deficits of variance in richness, the effect would not fluctuate between sites and years, as VNP admit it does. 3. The shape of the richness frequency distribution is a potentially interesting additional form of analysis, albeit one that we had chosen not to include in our analysis. However, it is surprising that VNP dismiss analysis of richness variance, but then interpret a more subtle aspect of the richness distribution — the skewness. VNP's redefinition of the terms‘Niche facilitation’for a deficit of low richness values, and ‘Niche limitation’ for a deficit of high values, leads them to misrepresent our arguments. They are mistaken in suggesting that niche limitation will necessarily lead to a skewed curve: they do not allow for the fact that the species frequencies, on which the null model is based already, incorporate effects of species interactions.‘Niche facilitation’, as defined by VNP, would lead to a variance excess, not a deficit as they assume. 4. VNP's criticism of a priori guild classifications had already been met by our use of the intrinsic guild approach. Guild analyses offer the best way forward. VNP use principally the methodology of 1987. There have been a number of methodological advances since that time. We used these advances in our original paper specifically to circumvent the kinds of problems that they identify. We agree with VNP that it would be very useful to explore the mechanisms behind assembly rules with experiments, but the logical first step is to identify potential assembly rules, as we have been attempting to do. We conclude that, whilst VNP make some interesting observations, none of their criticisms invalidate our results or conclusions. Our original approach stands as the best known approach, we believe, for searching for community structure in such data, and we reaffirm the validity of our ecological conclusions.     

Small-scale distribution of species richness in a grassland (Bílé Karpaty Mts., Czech Republic)

Variation in the number of species was studied in a subthermophilous grassland at a scale of 0.05 ×0.05 m during a 5-year period. The observed variance of species richness (VSR) was compared with a null model based on random distribution of species over a set of squares. It was found that distribution of species richness had more values than, expected around the mean and less values at the “shoulders”. Both tails fell within the predicted limits. Application of the procedures removing spatial dependence (random shifts, rotation/reflection method byPalmer & van der Maarel 1995) and environmental heterogeneity (patch model byWatkins & Wilson 1992) did not change the observed pattern. Using simulations in which the number of clumps and clumping intensity were manipulated it was found that the effect of the clumped spatial pattern, on VSR results in a wide range of variances. Both variance excess and variance deficit were found more frequently than expected under the null model. To test the effect of the limitation to the number of individuals per square, a null model was developed which included that observed number of plant shoots per square, the observed distribution of the number of shoots belonging to individual species per square and the observed spatial distribution of the shoots. The observed VSR was still lower than that produced by the null model. Therefore, it is concluded that at a scale of 0.05×0.05 m plant species combine in a non-random way in the studied grassland. It is suggested that the shape of left and right “shoulders” of the species richness distribution may be caused by different factors, such as positive and negative covariance between species, respectively. Their simultaneous impact can generate the observed pattern in species richness.     

Functional trait assembly through ecological and evolutionary time

A classic community assembly hypothesis is that all guilds must be represented before additional species from any given guild enter the community. We conceptually extend this hypothesis to continuous functional traits, refine the hypothesis with an eco-evolutionary model of interaction network community assembly, and compare the resultant continuous trait assembly rule to empirical data. Our extension of the “guild assembly rule” to continuous functional traits was rejected, in part, because the eco-evolutionary model predicted trait assembly to be characterized by the expansion of trait space and trait/species sorting within trait space. Hence, the guild rule may not be broadly applicable. A “revised” assembly rule did, however, emerge from the eco-evolutionary model: as communities assemble, the range in trait values will increase to a maximum and then remain relatively constant irrespective of further changes in species richness. This rule makes the corollary prediction that the trait range will, on average, be a saturating function of species richness. To determine if the assembly rule is at work in natural communities, we compared this corollary prediction to empirical data. Consistent with our assembly rule, trait “space” (broadly defined) commonly saturates with species richness. Our assembly rule may thus represent a general constraint placed on community assembly. In addition, taxonomic scale similarly influences the predicted and empirically observed relationship between trait “space” and richness. Empirical support for the model’s predictions suggests that studying continuous functional traits in the context of eco-evolutionary models is a powerful approach for elucidating general processes of community assembly.     

Contrasting patterns and drivers in taxonomic versus functional diversity,and community assembly of aquatic plants in subtropical lakes

Little is known about the differences in patterns and drivers between species richness (SR) and functional diversity (FD) in aquatic plants at large scales, and the underlying assembly mechanisms are not well studied. We compared SR and FD patterns of aquatic plant assemblages in 29 subtropical lakes, and detected the underlying assembly rules. Environmental drivers of SR and FD were revealed by GLM and GAM models, and the relative importance of assembly rules was determined by a null model approach. SR and FD of aquatic plants presented different patterns and drivers in this region. SR was significantly correlated with geographic, hydrological and water quality variables. We found a lower functional richness but higher functional evenness and divergence in the highland lakes. There was no significant correlation between functional richness and environmental variables. Null model analyses showed that most values of standardized effect size were located between the confidence interval, indicating a dominance of randomness. Deterministic processes such as limiting similarity and habitat filtering were also important in individual lakes. Habitat filtering plays a stronger role shaping the hydrophyte assemblages especially with the increase of elevation, area and AWLF (amplitude of water level fluctuation). Our results demonstrated that FD, in contrast to SR, were more resistant to environmental variations, and hydrology played an important role in shaping both SR and FD patterns in lake ecosystems. Furthermore, we revealed complex assembly rules and emphasized the importance of both stochastic and deterministic mechanisms in structuring aquatic plant assemblages at the regional scale.     

Ecological assembly rules in plant communities—approaches,patterns and prospects

Understanding how communities of living organisms assemble has been a central question in ecology since the early days of the discipline. Disentangling the different processes involved in community assembly is not only interesting in itself but also crucial for an understanding of how communities will behave under future environmental scenarios. The traditional concept of assembly rules reflects the notion that species do not co‐occur randomly but are restricted in their co‐occurrence by interspecific competition. This concept can be redefined in a more general framework where the co‐occurrence of species is a product of chance, historical patterns of speciation and migration, dispersal, abiotic environmental factors, and biotic interactions, with none of these processes being mutually exclusive. Here we present a survey and meta‐analyses of 59 papers that compare observed patterns in plant communities with null models simulating random patterns of species assembly. According to the type of data under study and the different methods that are applied to detect community assembly, we distinguish four main types of approach in the published literature: species co‐occurrence, niche limitation, guild proportionality and limiting similarity. Results from our meta‐analyses suggest that non‐random co‐occurrence of plant species is not a widespread phenomenon. However, whether this finding reflects the individualistic nature of plant communities or is caused by methodological shortcomings associated with the studies considered cannot be discerned from the available metadata. We advocate that more thorough surveys be conducted using a set of standardized methods to test for the existence of assembly rules in data sets spanning larger biological and geographical scales than have been considered until now. We underpin this general advice with guidelines that should be considered in future assembly rules research. This will enable us to draw more accurate and general conclusions about the non‐random aspect of assembly in plant communities.     

大型食草动物的聚群规则:印度西部热带干旱森林中有蹄类动物局域和区域尺度的多样性格局

竞争能够塑造自然群落并由其给出可用备择模型检验的一般性聚集规则。关于多种动物集合的竞争结构证据至今还非常少见,我们给出了一个备择模型分析用于检验印度西部热带干旱森林中当地5种大型食草类动物(花鹿Axisaxis,印度大蓝羚Boselaphustragocamelus,水鹿Cervusunicolor,印度瞪羚Gazellabennetti,野猪Susscrofa)是否符合一般性集合规则。使用蒙特卡罗模拟分析了动物的生态位重叠和身体大小比率的类似性,结果表明当地这些种的集合不是竞争结构决定的,其内部阶元结构处于亚最适种丰富度。另外2种有蹄类(印度黑羚Antilopecervicapra,四角羚羊Tetracerusquadricornis)属于地域性种库的组成部分,但缺失于这种集合;身体大小组合在地域性物种集合中不是随机的,然而在当地集合中能够通过随机性机会获得。对于观察到的类型,我们认为由于灭绝留下的统计学空缺似乎最能解释现存集合。由于天敌(大型食肉类)能够减少猎物物种的种间竞争,使得这种无规则生态位配置具有成立的可能性。由于对四角羚羊(T.quadricornis)的生物学和保护现状知之甚少,未将该种包括在该聚集中。我认为对这类较少被人类了解的物种其保护问题被忽视了,今后获得这些稀少的地方特有物种的生态学资料是一项紧急课题.     

Variability in species richness and guild structure in two species-rich grasslands

It has been suggested that variation in the proportion of species in guilds (=guild proportionality) indicates community structuring by guilds in biotic communities. This hypothesis was tested on a subthermophilous grassland and a mesotrophic meadow at a scale of 0.09 m² based on a five-year data set. Further, variation in the total number of species, variation in the number of species belonging to a guild and non-randomness in species composition of guilds were studied. A number of criteria for guild definition were used, such as life form, Grime's C-S-R strategy, phenology, plant height, pollination and dispersal syndromes, leaf shape and anatomy and taxonomy at the family level. The observed variation in the number of guild species corresponded to the null model in which species assemblages with fixed species richness per square were randomly generated from the species pool. The observed variation in the number of guild species was often higher than the variation calculated for randomly distributed species whereas the variation in the proportion of guild species was in some cases lower than the variation calculated for randomly distributed species with fixed frequencies. Possible reasons for the discrepancy in the results based on different models are discussed. It is concluded that there is little evidence of guilds in the organization of grasslands. *** DIRECT SUPPORT *** A02DO006 00012     

Native–Exotic Species Richness Relationships Across Spatial Scales in a Prairie Restoration Matrix

In highly invaded ecosystems, restoration of native plant communities is dependent upon reducing exotic species relative to native species. Even so, in monitoring, the native–exotic species richness ratio has been shown to be scale‐dependent. Measurement at small spatial scales (<1 m²) can reveal a negative native–exotic richness relationship, where niche occupation may prevent invasion. Conversely, at larger scales, a positive correlation may exist, where environmental heterogeneity and equally favorable conditions may drive native–exotic relationships. Here, we compare slopes of native–exotic relationships across spatial scales in a prairie undergoing active restoration. The observed native–exotic richness ratios varied considerably over scales ranging from 1 to 1,000 m², emphasizing the importance of choosing a measurement scale that is most pertinent to the treatment and ecological mechanism used to evaluate restoration success. Our native–exotic richness slopes were positive over all scales, but lower than would be expected in a random community assembly, suggesting the influence of niche‐based competition. Correspondingly, our native–exotic cover slope was more negative than a null model; however, areas of frequent fire treatments showed a significant deviation from null only for richness, indicating that burning may enhance native–exotic competitive dynamics for number of species but not cover. The negative native–exotic cover relationships appear to be driven in this system mainly by exotic graminoids, across burn treatments and native functional groups, supporting the concept that frequent burning can alter the dominant competitive mechanism from coverage of these exotic grasses to an improved environment for germination and dispersal of more native species.     

Testing for community structure: A bayesian approach

Variance deficit in richness is not reliable evidence for niche limitation. The main problem is the effect of limitation to individual plant module packing at small quadrat sizes, though environmental and historical patchiness can also confound the results unless patch models are used. More reliable approaches to community structure (assembly rules) are those that take into account the characters of the species—guild proportionality, texture convergence and limiting similarity test. A informally Bayesian approach to community structure is advocated, accepting that some hypotheses have greater prior probability, such as module-packing limitations, but using all the evidence available to estimate the likelihood of all the ecological hypotheses. The problem of environmental patchiness can be largely overcome by patch models. These models also overcome the problem of spatial autocorrelation, because they are conservative in such situations. For guild proportionality analyses, the possibility of sampling bias is a particular problem; this should be borne in mind during sampling, and checks made for the possibility in analyses. In devising Monte Carlo tests for community structure, permutation tests (i.e. randomisation tests, using sampling without replacement) are theoretically and practically preferable to bootstrap tests (i.e. using sampling with replacement). The best test is the simplest one that incorporates the intended null model and uses the intended test statistic. Experiments have intuitive appeal, but field experiments have several severe drawbacks, including the inevitability of artefacts, and insoluble problems of when to record. In practice, field experiments have told us surprisingly little about community structure.     

Limitations to species coexistence: evidence for competition from field observations,using a patch model

Abstract. Plant community structure in four dune slacks is examined for evidence of competition, expressed as local constraint on species biomass. Such constraint would result in a low variance in total quadrat biomass, compared to a null model. A method of analysis is introduced, using a patchbased null model, which is intended to discount most of the effects of environmental heterogeneity. The method is applied to data in which previous methods had failed to find such structure. There was significant biomass constraint (i.e. low variance in biomass) examining total plant-community biomass in Site 1, and there was a trend in this direction also in Site 2. When guilds defined on morphology were examined, significant biomass constraint was found for the Rhizomatous guild in Site 2. There was a trend to low variance for this guild also in Site 4, but it was significant only in a one-tailed test. After a consideration of other mechanisms, it is concluded that the most likely explanation for this biomass constraint is interspecies competition. Site 2, which showed the strongest evidence of biomass constraint in the present analyses, was also the site that demonstrated a significant deficit of variance in richness in earlier work, and was the most species-rich of the four sites and the least disturbed. The advantages of the new method, for obtaining evidence on competition, are discussed in comparison to pot experiments and field-perturbation experiments. We conclude that community structure can be seen in the pattern of species biomasses provided that a suitable method of analysis is used, i.e. examining small-scale effects and excluding the effects of environmental heterogeneity. In some cases such effects can be related to guild membership. We hypothesise that constraint on biomass is likely to be stronger in more species-rich and less disturbed communities.     

Change in pattern diversity during secondary succession in Estonian forests

Comparisons of observed variance in species diversity (exp LT) and in species richness with expectation assuming a random and independent distribution of species are used to assess the relative importance during succession of niche limitation (Wilson, Gitay & Agnew 1987; limitation of the abundance or occurrence of species by competitive interactions) and nucleation (Yarranton & Morrison 1974; the development of a community through chance establishment or persistence followed by vegetative expansion from those nuclei). Wilson, Gitay & Agnew (1987) and Palmer (1987) suggest a deficit (i.e. lower than expected) in variance of richness is consistent with a ‘niche limitation’ process where competitive sorting makes patches more similar in species number than would be expected. In contrast, variance that is greater than expected is consistent with and could result from underlying environmental heterogeneity (i.e. ‘waterhole effect’). We extend these ideas to suggest that, in addition, a deficit in variance of diversity (expLT) can be interpreted as indicating niche limitation. Further, we suggest that a deficit of variance in richness need not be interpreted as resulting from niche limitation, but could result from limited dispersal and establishment followed by localized spread that causes quadrats to have less compositional overlap than might be expected if species were distributed at random. However, there is little reason to expect that such nucleation would cause a similar reduction in the variance of diversity. First, expLT is relatively insensitive to the presence or absence of rare species, and second, if local competitive processes do restrict the values of exp LT through some form of niche limitation, this could well be independent of the specific species involved. Thus, a combination of lower than expected variance in richness and high variance in diversity suggests nucleation to be important, particularly if the mean pairwise similarity between samples is lower than expected. We first examine a secondary successional sere with stands representing post-logging and mature forest. The variance of diversity is low in the establishment phase, but not in subsequent phases where the tree canopy is well developed. Significantly low variance of species richness is observed in a young forest just entering the thinning phase. Thus, nucleation may be taking place in the establishment phase of forest development, and is almost certainly important in the early thinning phase. Second, we examine communities in the process of equilibration following fertilization, logging, and paludification. All these communities have a low variance in diversity (exp LT) suggesting niche limitation.     

Body-size differences in a colonizing amphipod-mollusc assemblage

Summary We examined body size patterns in a colonizing assemblage of marine amphipods and molluscs. We collected animals over a 25 day period from an archipelago of pits that were drilled in brick surfaces. The percent of pits occupied, abundance, species richness, and body size of colonists all increased significantly through time. We compared size ratios of coexisting species with two null models, one that randomized individuals and one that randomized species-populations. For both models, observed overlap ratios usually did not differ from randomness, although species richness was consistently lower than expected for later samples. Results were similar for a subset of the data, a guild of suspension/deposit feeding amphipod species. Some assemblages did show significant deviations from the null models, but the results were spotty and varied among replicates. Overall, the evidence for non-random overlap ratios in body sizes of colonizers was weak.     

Book reviews

The variance test, originally proposed for testing species associations, is used to test whether species richness is more or less variable than expected under the null model of no species interactions. Species richness is more variable than expected in some fields, and less variable than expected in other fields at the Cedar Creek Natural History Area in Minnesota. High variance in species richness may be caused by variability in competitive exclusion rates or small-scale environmental heterogeneity. Low variance in species richess may occur if the community is saturated with species, or if species-rich areas have high local competitive exclusion rates. Results of the variance test depend somewhat on quadrat size, and can be used to select study sites and quadrat sizes for further research on the nature of variation in species richness.     

Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain,Southwest China

Characterizing trait variation across different ecological scales in plant communities has been viewed as a way to gain insights into the mechanisms driving species coexistence. However, little is known about how changes in intraspecific and interspecific traits across sites influence species richness and community assembly, especially in understory herbaceous communities. Here we partitioned the variance of four functional traits (maximum height, leaf thickness, leaf area and specific leaf area) across four nested biological scales: individual, species, plot, and elevation to quantify the scale-dependent distributions of understory herbaceous trait variance. We also integrated the comparison of the trait variance ratios to null models to investigate the effects of different ecological processes on community assembly and functional diversity along a 1200-m elevational gradient in Yulong Mountain. We found interspecific trait variation was the main trait variation component for leaf traits, although intraspecific trait variation ranged from 10% to 28% of total variation. In particular, maximum height exhibited high plasticity, and intraspecific variation accounted for 44% of the total variation. Despite the fact that species composition varied across elevation and species richness decreased dramatically along the elevational gradient, there was little variance at our largest (elevation) scale in leaf traits and functional diversity remained constant along the elevational gradient, indicating that traits responded to smaller scale influences. External filtering was only observed at high elevations. However, strong internal filtering was detected along the entire elevational gradient in understory herbaceous communities, possibly due to competition. Our results provide evidence that species coexistence in understory herbaceous communities might be structured by differential niche-assembled processes. This approach--integrating different biological scales of trait variation--may provide a better understanding of the mechanisms involved in the structure of communities.