Advances, challenges and a developing synthesis of ecological community assembly theory

Ecological approaches to community assembly have emphasized the interplay between neutral processes, niche-based environmental filtering and niche-based species sorting in an interactive milieu. Recently, progress has been made in terms of aligning our vocabulary with conceptual advances, assessing how trait-based community functional parameters differ from neutral expectation and assessing how traits vary along environmental gradients. Experiments have confirmed the influence of these processes on assembly and have addressed the role of dispersal in shaping local assemblages. Community phylogenetics has forged common ground between ecologists and biogeographers, but it is not a proxy for trait-based approaches. Community assembly theory is in need of a comparative synthesis that addresses how the relative importance of niche and neutral processes varies among taxa, along environmental gradients, and across scales. Towards that goal, we suggest a set of traits that probably confer increasing community neutrality and regionality and review the influences of stress, disturbance and scale on the importance of niche assembly. We advocate increasing the complexity of experiments in order to assess the relative importance of multiple processes. As an example, we provide evidence that dispersal, niche processes and trait interdependencies have about equal influence on trait-based assembly in an experimental grassland.     

A trait-based approach to community assembly: partitioning of species trait values into within- and among-community components

Plant functional traits vary both along environmental gradients and among species occupying similar conditions, creating a challenge for the synthesis of functional and community ecology. We present a trait-based approach that provides an additive decomposition of species' trait values into alpha and beta components: beta values refer to a species' position along a gradient defined by community-level mean trait values; alpha values are the difference between a species' trait values and the mean of co-occurring taxa. In woody plant communities of coastal California, beta trait values for specific leaf area, leaf size, wood density and maximum height all covary strongly, reflecting species distributions across a gradient of soil moisture availability. Alpha values, on the other hand, are generally not significantly correlated, suggesting several independent axes of differentiation within communities. This trait-based framework provides a novel approach to integrate functional ecology and gradient analysis with community ecology and coexistence theory.     

Functional traits reveal the expansion and packing of ecological niche space underlying an elevational diversity gradient in passerine birds

Variation in species richness across environmental gradients may be associated with an expanded volume or increased packing of ecological niche space. However, the relative importance of these alternative scenarios remains unknown, largely because standardized information on functional traits and their ecological relevance is lacking for major diversity gradients. Here, we combine data on morphological and ecological traits for 523 species of passerine birds distributed across an Andes-to-Amazon elevation gradient. We show that morphological traits capture substantial variation in species dietary (75%) and foraging niches (60%) when multiple independent trait dimensions are considered. Having established these relationships, we show that the 14-fold increase in species richness towards the lowlands is associated with both an increased volume and density of functional trait space. However, we find that increases in volume contribute little to changes in richness, with most (78%) lowland species occurring within the range of trait space occupied at high elevations. Taken together, our results suggest that high species richness is mainly associated with a denser occupation of functional trait space, implying an increased specialization or overlap of ecological niches, and supporting the view that niche packing is the dominant trend underlying gradients of increasing biodiversity towards the lowland tropics.     

A trait-based approach predicting community assembly and dominance of microbial invasive species

Understanding the mechanisms underlying community assembly helps to define success and susceptibility to biological invasions. Here, we explored phytoplankton community assembly following niche and neutral paradigms and using a trait-based approach. Under the hypothesis that the morphology-based functional groups (MBFG) clusters species with similar niche, we analysed how trait-related differences in fitness influence dominance of an invasive species. This was based on literature review, field data and model simulations. We predict that invading species can be dominant if: 1) do not belong to the local MBFG but use unexploited areas of the niche, or 2) belong to the resident MBFG but exhibit a higher fitness due to a particular combination of traits. The invasive dinoflagellate Ceratium furcoides was used as the model species to evaluate these hypotheses, its morphological (e.g. volume) and physiological (e.g. growth rates) traits were compared with species from the same (V: photosynthetic flagellates) and different (VII: colonial cyanobacteria) MBFG. Fitness was estimated using models parametrized with MBFG rates (R*, ability to draw down phosphate) under different environmental conditions (i.e. flushing). Results contributed to support both hypotheses. First, the alternation of C. furcoides and cyanobacteria dominance was explained by the use of different niches. Secondly, species from MBFG V were dominant under similar environments. Within this group V C. furcoides showed higher fitness under low flushing and high predation, advantage provided by a distinctive combination of traits. The application of trait-based approaches to represent the niche and estimate fitness along environmental gradients was useful to evaluate community assembly and can be used to predict the dominance of microbial species invasions.     

Intraspecific trait variation across elevation predicts a widespread tree species' climate niche and range limits

Global change is widely altering environmental conditions which makes accurately predicting species range limits across natural landscapes critical for conservation and management decisions. If climate pressures along elevation gradients influence the distribution of phenotypic and genetic variation of plant functional traits, then such trait variation may be informative of the selective mechanisms and adaptations that help define climatic niche limits. Using extensive field surveys along 16 elevation transects and a large common garden experiment, we tested whether functional trait variation could predict the climatic niche of a widespread tree species (Populus angustifolia) with a double quantile regression approach. We show that intraspecific variation in plant size, growth, and leaf morphology corresponds with the species' total climate range and certain climatic limits related to temperature and moisture extremes. Moreover, we find evidence of genetic clines and phenotypic plasticity at environmental boundaries, which we use to create geographic predictions of trait variation and maximum values due to climatic constraints across the western US. Overall, our findings show the utility of double quantile regressions for connecting species distributions and climate gradients through trait‐based mechanisms. We highlight how new approaches like ours that incorporate genetic variation in functional traits and their response to climate gradients will lead to a better understanding of plant distributions as well as identifying populations anticipated to be maladapted to future environments.     

Trait plasticity,not values,best corresponds with woodland plant success in novel and manipulated habitats

Aims The clustering of plants with similar leaf traits along environmental gradients may arise from adaptation as well as acclimation to heterogeneous habitat conditions. Determining the forces that shape plant leaf traits requires both linking variation in trait morphology with abiotic gradients and linking that trait variation with plant performance under varying abiotic conditions. Across the spectrum of plant types, shade-tolerant evergreen herbs are relatively low in trait plasticity, compared to deciduous and sun-adapted species. These plants employ stress-tolerant strategies for survival, which coincide with relatively static trait morphologies, slow growth and hence a lower ability to adjust to changing environmental conditions.Methods We investigate how the survival of two ecologically similar understory evergreen species, Asarum arifolium and Hepatica nobilis, corresponds with variation in six commonly measured functional traits (leaf area, specific leaf area, plant height, leaf number, leaf length and shoot mass) along natural and experimental abiotic gradients. We examine temporal (the period 2007–9) and spatial (100 km) variations in these traits after (i) translocating 576 plants across a span from the southern Appalachian Mountains in NC, USA, to the Piedmont, GA, USA, which includes north- and south-facing slope habitats and (ii) the experimental manipulation of diffuse light and soil moisture.Important findings We find that when translocated into a novel habitats, with novel environmental conditions that often are more extreme than the source habitat, both species appear capable of considerable morphological acclimation and generally converge to similar trait values. Hepatica nobilis does not exhibit mean trait values particularly different from those of A. arifolium, but it demonstrates much greater phenotypic plasticity. These results indicate that relatively conservative plant species nonetheless acclimate and survive across heterogeneous environmental conditions.     

Joint effect of phylogenetic relatedness and trait selection on the elevational distribution of Rhododendron species

Congeneric species may coexist at fine spatial scales through niche differentiation, however, the magnitude to which the effects of functional traits and phylogenetic relatedness contribute to their distribution along elevational gradients remains understudied. To test the hypothesis that trait and elevational range overlap can affect local speciesʼ coexistence, we first compared phylogenetic relatedness and trait (including morphological traits and leaf elements) divergence among closely related species of Rhododendron L. on Yulong Mountain, China. We then assessed relationships between the overlap of multiple functional traits and the degree of elevational range overlap among species pairs in a phylogenetic context. We found that phylogeny was a good predictor for most functional traits, where closely related species showed higher trait similarity and occupied different elevational niches at our study site. Species pairs of R. subgen. Hymenanthes (Blume) K. Koch showed low elevational range overlap and some species pairs of R. subgen. Rhododendron showed obvious niche differentiation. Trait divergence is greater for species in R. subgen. Rhododendron, and it plays an important role between species pairs with low elevational range overlap. Trait convergent selection takes place between co-occurring closely related species that have high elevational range overlap, which share more functional trait space due to environmental filtering or ecological adaptation in more extreme habitats. Our results highlight the importance of evolutionary history and trait selection for species coexistence at fine ecological scales along environmental gradients.     

Contrasting Effects of Intraspecific Trait Variation on Trait-Based Niches and Performance of Legumes in Plant Mixtures

Niche differentiation, assumed to be a key mechanism of species coexistence, requires that species differ in their functional traits. So far it remains unclear to which extent trait plasticity leads to niche shifts of species at higher plant diversity, thereby increasing or decreasing niche overlap between species. To analyse this question it is convenient to measure niches indirectly via the variation in resource-uptake traits rather than directly via the resources used. We provisionally call these indirectly measured niches trait-based niches. We studied shoot- and leaf-morphological characteristics in seven legume species in monoculture and multi-species mixture in experimental grassland. Legume species varied in the extent of trait variation in response to plant diversity. Trait plasticity led to significant shifts in species niches in multiple dimensions. Single-species niches in several traits associated with height growth and filling of canopy space were expanded, while other niche dimensions were compressed or did not change with plant diversity. Niche separation among legumes decreased in dimensions related to height growth and space filling, but increased in dimensions related to leaf size and morphology. The total extent of occupied niche space was larger in mixture than in the combined monocultures for dimensions related to leaf morphology and smaller for dimensions related to whole-plant architecture. Taller growth, greater space filling and greater plasticity in shoot height were positively, while larger values and greater plasticity in specific leaf area were negatively related with increased performance of species in mixture. Our study shows that trait variation in response to plant diversity shifts species niches along trait axes. Plastically increased niche differentiation is restricted to niche dimensions that are apparently not related to size-dependent differences between species, but functional equivalence (convergence in height growth) rather than complementarity (divergence in traits associated with light acquisition) explains increased performance of legumes in mixture.     

Trait‐based community assembly of aquatic macrophytes along a water depth gradient in a freshwater lake

相似文献   

Functional traits of tree species with phylogenetic signal co-vary with environmental niches in two large forest dynamics plots

Aims While using phylogenetic and functional approaches to test the mechanisms of community assembly, functional traits often act as the proxy of niches. However, there is little detailed knowledge regarding the correlation between functional traits of tree species and their niches in local communities. We suggest that the co-varying correlation between functional traits and niches should be the premise for using phylogenetic and functional approaches to test mechanisms of community assembly. Using functional traits, phylogenetic and environmental data, this study aims to answer the questions: (i) within local communities, do functional traits of co-occurring species co-vary with their environmental niches at the species level? and (ii) what is the key ecological process underlying community assembly in Xishuangbanna and Ailaoshan forest dynamic plots (FDPs)?Methods We measured seven functional traits of 229 and 36 common species in Xishuangbanna and Ailaoshan FDPs in tropical and subtropical China, respectively. We also quantified the environmental niches for these species based on conditional probability. We then analyzed the correlations between functional traits and environmental niches using phylogenetic independent contrasts. After examining phylogenetic signals of functional traits using Pagel's λ, we quantified the phylogenetic and functional dispersion along environmental gradients within local tree communities.Important findings For target species, functional traits do co-vary with environmental niches at the species level in both of the FDPs, supporting that functional traits can be used as a proxy for local-scale environmental niches. Functional traits show significant phylogenetic signals in both of the FDPs. We found that the phylogenetic and functional dispersion were significantly clustered along topographical gradients in the Ailaoshan FDP but overdispersion in the Xishuangbanna FDP. These patterns of phylogenetic and functional dispersion suggest that environmental filtering plays a key role in structuring local tree assemblages in Ailaoshan FDP, while competition exclusion plays a key role in Xishuangbanna FDP.     

Trait plasticity in species interactions: a driving force of community dynamics

Evolutionary community ecology is an emerging field of study that includes evolutionary principles such as individual trait variation and plasticity of traits to provide a more mechanistic insight as to how species diversity is maintained and community processes are shaped across time and space. In this review we explore phenotypic plasticity in functional traits and its consequences at the community level. We argue that resource requirement and resource uptake are plastic traits that can alter fundamental and realised niches of species in the community if environmental conditions change. We conceptually add to niche models by including phenotypic plasticity in traits involved in resource allocation under stress. Two qualitative predictions that we derive are: (1) plasticity in resource requirement induced by availability of resources enlarges the fundamental niche of species and causes a reduction of vacant niches for other species and (2) plasticity in the proportional resource uptake results in expansion of the realized niche, causing a reduction in the possibility for coexistence with other species. We illustrate these predictions with data on the competitive impact of invasive species. Furthermore, we review the quickly increasing number of empirical studies on evolutionary community ecology and demonstrate the impact of phenotypic plasticity on community composition. Among others, we give examples that show that differences in the level of phenotypic plasticity can disrupt species interactions when environmental conditions change, due to effects on realized niches. Finally, we indicate several promising directions for future phenotypic plasticity research in a community context. We need an integrative, trait-based approach that has its roots in community and evolutionary ecology in order to face fast changing environmental conditions such as global warming and urbanization that pose ecological as well as evolutionary challenges.     

Complex patterns of environmental niche evolution in Iberian columbines (genus Aquilegia,Ranunculaceae)

Aims This study explores the patterns of niche differentiation in a group of seven closely related columbines (genus Aquilegia, Ranunculaceae) from the Iberian Peninsula. Populations of these columbines are subject to complex patterns of divergent selection across environments, which partly explain the taxonomic structure of the group. This suggests the hypothesis that niche divergence must have occurred along the process of diversification of the group.Methods We used MaxEnt to build environmental niche models of seven subspecies belonging to the three species of Aquilegia present in the Iberian Peninsula. From these models, we compared the environmental niches through two different approaches: ENMtools and multivariate methods.Important findings MaxEnt distributions conformed closely to the actual distribution of the study taxa. ENMtools methods failed to uncover any clear patterns of niche differentiation or conservatism in Iberian columbines. Multivariate analyses indicate the existence of differentiation along altitudinal gradients and along a gradient of climatic conditions determined by the summer precipitation and temperatures. However, climatic conditions related to winter temperature and precipitation, as well as soil properties, were equally likely to show conservatism or divergence. The complex patterns of niche evolution we found suggest that Iberian Columbines have not been significantly constrained by forces of niche conservatism, so they could respond adaptively to the fast and profound climate changes in the Iberian Peninsula through the glacial cycles of the Pleistocene.     

Stress from cold and drought as drivers of functional trait spectra in North American angiosperm tree assemblages

Understanding how environmental change alters the composition of plant assemblages, and how this in turn affects ecosystem functioning is a major challenge in the face of global climate change. Assuming that values of plant traits express species adaptations to the environment, the trait‐based approach is a promising way to achieve this goal. Nevertheless, how functional traits are related to species’ environmental tolerances and how trait spectra respond to broad‐scale environmental gradients remains largely unexplored. Here, we identify the main trait spectra for US angiosperm trees by testing hypotheses for the relationships between functional traits and species’ environmental tolerances to environmental stresses, as well as quantifying the environmental drivers of assemblage means and variances of these traits. We analyzed >74,000 community assemblages from the US Forest Inventory and Analysis using 12 functional traits, five traits expressing species’ environmental tolerances and 10 environmental variables. Results indicated that leaf traits, dispersal traits, and traits related to stem hydraulics were related to cold or drought tolerance, and their assemblage means were best explained by minimum temperatures. Assemblage means of traits related to shade tolerance (tree growth rate, leaf phosphorus content, and bark thickness) were best explained by aridity index. Surprisingly, aridity index, rather than minimum temperature, was the best predictors of assemblage variances of most traits, although these relationships were variable and weak overall. We conclude that temperature is likely to be the most important driver of functional community structure of North American angiosperm trees by selecting for optimum strategies along the cold and drought stress trade‐off. In turn, water availability primarily affects traits related to shade tolerance through its effect on forest canopy structure and vegetation openness.     

A trait-based approach across the native and invaded range to understand plant invasiveness and community impact

Alien plant species invasiveness and impact on diversity (i.e. species richness and composition) can be driven by the altered competitive interactions experienced by the invader in its invaded range compared to its native range. Trait-based competition effects on invasiveness can be mediated through size-asymmetric competition, i.e. a trait suit of the invader that drives competitive dominance, and through ‘niche differences', i.e. trait differentiation and thus minimized competition between invader and the invaded community. In terms of invasion impact, size-asymmetric competition is expected to result in competitive exclusion of co-occurring subordinate species, whereas ‘niche differences' might result in competitive exclusion of the most functionally similar co-occurring species. Although observational work does not allow the full disentanglement of both trait-based effects, it does allow to verify the occurrence of expected theoretical trait patters. In this study, we explored the trait-based competition effects on invasiveness and diversity impact for Rosa rugosa in both its invaded range in Belgium and its native range in Japan, based on seven functional traits across 100 vegetation plots. Following the predictions for enhanced invasiveness, we found much lower functional overlap between R. rugosa and the co-occurring species in the invaded range compared to the native range. This likely also explains the absence of diversity impact in its native range. Despite the absence of changes in species richness in the invaded range, the invader did strongly impact species composition of invaded communities. This impact occurred through strong shade tolerance responses, suggesting size-asymmetric competition effects and cover loss of co-occurring dominant species, next to exclusion of co-occurring species most functionally similar to the invader; suggesting niche difference effects. In conclusion, this case-study illustrates how exploring functional trait patterns across a species native and invaded range can help in understanding how trait-based competition processes can affect invasiveness and community impact.     

Coupled plant traits adapted to wetting/drying cycles of substrates co-define niche multidimensionality

Theories attempting to explain species coexistence in plant communities have argued in favour of species' capacities to occupy a multidimensional niche with spatial, temporal and biotic axes. We used the concept of hydrological niche segregation to learn how ecological niches are structured both spatially and temporally and whether small scale humidity gradients between adjacent niches are the main factor explaining water partitioning among tree species in a highly water-limited semiarid forest ecosystem. By combining geophysical methods, isotopic ecology, plant ecophysiology and anatomical measurements, we show how coexisting pine and oak species share, use and temporally switch between diverse spatially distinct niches by employing a set of functionally coupled plant traits in response to changing environmental signals. We identified four geospatial niches that turned into nine, when considering the temporal dynamics of the wetting/drying cycles in the substrate and the particular plant species adaptations to garner, transfer, store and use water. Under water scarcity, pine and oak exhibited water use segregation from different niches, yet under maximum drought when oak trees crossed physiological thresholds, niche overlap occurred. The identification of niches and mechanistic understanding of when and how species use them will help unify theories of plant coexistence and competition.     

Quantifying the role of deterministic assembly and stochastic drift in a natural community of Arctic mosses

The interpretation of natural plant communities frequently invokes species‐sorting controlled by niche differences along spatial environmental gradients. This process of niche structuring can be explained by reference to functional traits, which provide a mechanistic explanation for community structure. In contrast, models explaining species coexistence obviate the limiting effect of niche difference, by invoking processes which cause species‐level drift, e.g. demographic stochasticity. This paper investigates a simple habitat with strong gradients (moss communities in a patterned arctic wetland) to identify signature‐patterns under‐pinning the relative importance of deterministic assembly and stochastic drift in a natural community. First, ordination analysis was used to confirm community composition structured by a range of nine carefully selected functional traits. Second, to determine whether traits explaining community composition might also explain species richness, local species richness (sR) was compared to (1) observed trait diversity and (2) expected trait diversity based on permutation tests, which are used to simulate null community assembly for different values of sR. Traits explaining species composition, consistent with deterministic niche structuring, do not appear to maintain sR. This surprising result was explained by decomposing the community into individual pair‐wise comparisons, i.e. species niche‐differences and association (χ²). Results support deterministic processes via the sorting of species with similar and contrasting niches, at opposite ends of a composite environmental gradient. Nevertheless, stochastic drift is apparent in the random structure of a majority of pair‐wise associations; in addition, a species’ abundance was in general not related to environmental distance from response‐optima. We suggest therefore that spatial pattern in the moss community is a balance between deterministic forces with respect to species traits and controlling environmental gradients, and stochastic drift, which weakens this deterministic structure.     

Plant functional and phylogenetic turnover correlate with climate and land use in the Western Swiss Alps

Aims Understanding the relative importance of historical and environmental processes in the structure and composition of communities is one of the longest quests in ecological research. Increasingly, researchers are relying on the functional and phylogenetic β-diversity of natural communities to provide concise explanations on the mechanistic basis of community assembly and the drivers of trait variation among species. The present study investigated how plant functional and phylogenetic β-diversity change along key environmental and spatial gradients in the Western Swiss Alps.Methods Using the quadratic diversity measure based on six functional traits—specific leaf area, leaf dry matter content, plant height, leaf carbon content, leaf nitrogen content and leaf carbon to nitrogen content alongside a species-resolved phylogenetic tree—we relate variations in climate, spatial geographic, land use and soil gradients to plant functional and phylogenetic turnover in mountain communities of the Western Swiss Alps.Important findings Our study highlights two main points. First, climate and land-use factors play an important role in mountain plant community turnover. Second, the overlap between plant functional and phylogenetic turnover along these gradients correlates with the low phylogenetic signal in traits, suggesting that in mountain landscapes, trait lability is likely an important factor in driving plant community assembly. Overall, we demonstrate the importance of climate and land-use factors in plant functional and phylogenetic community turnover and provide valuable complementary insights into understanding patterns of β-diversity along several ecological gradients.     

Incorporating intraspecific variation in tests of trait-based community assembly

Environmental filtering and niche differentiation are processes proposed to drive community assembly, generating nonrandom patterns in community trait distributions. Despite the substantial intraspecific trait variation present in plant communities, most previous studies of trait-based community assembly have used species mean trait values and therefore not accounted for intraspecific variation. Using a null model approach, I tested for environmental filtering and niche differentiation acting on three key functional traits??vegetative height, specific leaf area (SLA), and leaf dry matter content (LDMC)??in old-field plant communities. I also examined how accounting for intraspecific variation at the among-plot and individual levels affected the detection of nonrandom assembly patterns. Tests using fixed species mean trait values provided evidence of environmental filtering acting on height and SLA and niche differentiation acting on SLA. Including plot-level intraspecific variation increased the strength of these patterns, indicating an important role of intraspecific variation in community assembly. Tests using individual trait data indicated strong environmental filtering acting on all traits, but provided no evidence of niche differentiation, although these signals may have been obscured by the effects of dispersal limitation and spatial aggregation of conspecific individuals. There was also strong evidence of nonrandom assembly of individuals within single species, with the strength of environmental filtering varying among species. This study demonstrates that, while analyses using fixed species mean trait values can provide insights into community assembly processes, accounting for intraspecific variation provides a more complete view of communities and the processes driving their assembly.     

Functional diversity of crustacean zooplankton communities: towards a trait-based classification

1. While studies of phytoplankton and terrestrial plant communities have increasingly emphasised the use of functional traits in ecological research, few have yet to apply this approach to zooplankton communities.
2. This study reviews laboratory and observational studies on zooplankton feeding and life history and provides a series of functional trait tables for the North American freshwater zooplankton. Qualitative and quantitative trait tables highlight areas where data were more scarce and point to which types of studies could fill in gaps in our knowledge of zooplankton niches.
3. Data were most complete for the Cladocera across most traits, while feeding information for cyclopoids was most sparse. Qualitative data that distinguished congeneric species were lacking for most groups.
4. A regional community dendrogram for common north-eastern North American zooplankton species was generated and shows that taxonomic differences between species do not capture fully functional differences based on the traits of body length, habitat, trophic group and feeding type.
5. The data collected here, combined with readily measurable species attributes, can be used to generate a multivariate measure of the functional niche of each species found in a community. Armed with this information, functional relationships that are useful for ecological studies of lake ecosystems can be more easily conducted.     

Functional traits,convergent evolution,and periodic tables of niches

Ecology is often said to lack general theories sufficiently predictive for applications. Here, we examine the concept of a periodic table of niches and feasibility of niche classification schemes from functional trait and performance data. Niche differences and their influence on ecological patterns and processes could be revealed effectively by first performing data reduction/ordination analyses separately on matrices of trait and performance data compiled according to logical associations with five basic niche ‘dimensions’, or aspects: habitat, life history, trophic, defence and metabolic. Resultant patterns then are integrated to produce interpretable niche gradients, ordinations and classifications. Degree of scheme periodicity would depend on degrees of niche conservatism and convergence causing species clustering across multiple niche dimensions. We analysed a sample data set containing trait and performance data to contrast two approaches for producing niche schemes: species ordination within niche gradient space, and niche categorisation according to trait‐value thresholds. Creation of niche schemes useful for advancing ecological knowledge and its applications will depend on research that produces functional trait and performance datasets directly related to niche dimensions along with criteria for data standardisation and quality. As larger databases are compiled, opportunities will emerge to explore new methods for data reduction, ordination and classification.