Sensitivity of plant functional types to climate change: classification tree analysis of a simulation model

Question: The majority of studies investigating the impact of climate change on local plant communities ignores changes in regional processes, such as immigration from the regional seed pool. Here we explore: (i) the potential impact of climate change on composition of the regional seed pool, (ii) the influence of changes in climate and in the regional seed pool on local community structure, and (iii) the combinations of life history traits, i.e. plant functional types (PFTs), that are most affected by environmental changes. Location: Fire‐prone, Mediterranean‐type shrublands in southwestern Australia. Methods: Spatially explicit simulation experiments were conducted at the population level under different rainfall and fire regime scenarios to determine the effect of environmental change on the regional seed pool for 38 PFTs. The effects of environmental and seed immigration changes on local community dynamics were then derived from community‐level experiments. Classification tree analyses were used to investigate PFT‐specific vulnerabilities to climate change. Results: The classification tree analyses revealed that responses of PFTs to climate change are determined by specific trait characteristics. PFT‐specific seed production and community patterns responded in a complex manner to climate change. For example, an increase in annual rainfall caused an increase in numbers of dispersed seeds for some PFTs, but decreased PFT diversity in the community. Conversely, a simulated decrease in rainfall reduced the number of dispersed seeds and diversity of PFTs. Conclusions: PFT interactions and regional processes must be considered when assessing how local community structure will be affected by environmental change.     

An improved method for searching plant functional types by numerical analysis

Abstract. The use of plant functional types (PFTs) to describe patterns and processes in plant communities has become essential to study and predict consequences of global change on vegetation and ecosystem processes. A PFT is a group of plants that, irrespective of phylogeny, are similar in a given set of traits and similar in their association to certain variables, which may be factors to which the plants are responding or effects of the plants in the ecosystem. To define PFTs relevant traits must be selected and an appropriate method must be used to classify plants into types. We critically review methods used for the analysis of PFT‐based data and describe a new recursive algorithm to numerically search for traits and find optimal PFTs. The algorithm uses three data matrices: describing populations by traits, communities by these populations and community sites by environmental factors or effects. It defines PFTs polythetically by cluster analysis, revealing plant types whose performance in communities is maximally associated to the specified environmental variables. We test the method with data from natural grassland communities of southern Brazil, which were experimentally subjected to combinations of grazing levels and N‐fertilizer. The new method is found to be better than similar analytical procedures previously described. Redundancy among traits is discussed and a procedure for comparing alternative solutions is presented based on the similarity in terms of PFT responses between different trait subsets. The concept of PFT response group is illustrated by example.     

A functional trait-based approach to understand community assembly and diversity–productivity relationships over 7 years in experimental grasslands

Several multi-year biodiversity experiments have shown positive species richness–productivity relationships which strengthen over time, but the mechanisms which control productivity are not well understood. We used experimental grasslands (Jena Experiment) with mixtures containing different numbers of species (4, 8, 16 and 60) and plant functional groups (1–4; grasses, legumes, small herbs, tall herbs) to explore patterns of variation in functional trait composition as well as climatic variables as predictors for community biomass production across several years (from 2003 to 2009). Over this time span, high community mean trait values shifted from the dominance of trait values associated with fast growth to trait values suggesting a conservation of growth-related resources and successful reproduction. Increasing between-community convergence in means of several productivity-related traits indicated that environmental filtering and exclusion of competitively weaker species played a role during community assembly. A general trend for increasing functional trait diversity within and convergence among communities suggested niche differentiation through limiting similarity in the longer term and that similar mechanisms operated in communities sown with different diversity. Community biomass production was primarily explained by a few key mean traits (tall growth, large seed mass and leaf nitrogen concentration) and to a smaller extent by functional diversity in nitrogen acquisition strategies, functional richness in multiple traits and functional evenness in light-acquisition traits. Increasing species richness, presence of an exceptionally productive legume species (Onobrychis viciifolia) and climatic variables explained an additional proportion of variation in community biomass. In general, community biomass production decreased through time, but communities with higher functional richness in multiple traits had high productivities over several years. Our results suggest that assembly processes within communities with an artificially maintained species composition maximize functional diversity through niche differentiation and exclusion of weaker competitors, thereby maintaining their potential for high productivity.     

Plant species’ coexistence relationships may shift according to life history traits and seasons

Although several studies have demonstrated that disturbance contributes to species’ diversity, little emphasis has been placed on the identification of species’ coexistence mechanisms related to life history traits. In this study, we compared species’ richness and components of plant communities around river confluences to explore how disturbance promotes the coexistence of species with different life history traits. Sites upstream and downstream of confluences are ideal for such comparisons because they draw on the same species’ pools and have similar ambient conditions, but differ markedly in the extents of flooding disturbance. We compared sites upstream and downstream of confluences by calculating species’ richness and community similarity indices for several life history traits in both summer and spring. In summer, the combined richness of all the species, of annual- and summer-flowering species, was higher downstream from confluences than upstream, but this was not the case for perennials. Similarity analyses suggested that plant communities are constructed according to a neutral process, whereby interactions between the coexisting species are neutral. However, in spring, species’ richness was similar upstream and downstream of confluences for all life history traits. Similarity analyses suggested that under these circumstances, the communities were constructed through a species-sorting process; i.e., each life history trait had a distinct habitat preference. Thus, the relative strengths of different community assembly processes may change seasonally. We concluded that species groups differing in their responses to disturbance may coexist in a single community. Thus, community structuring following disturbance may involve two processes: a neutral and a species-sorting process. The relative importance of each may vary between species’ life history traits and between seasons, and the interaction may account for current community structures.     

Constraints on trait combinations explain climatic drivers of biodiversity: the importance of trait covariance in community assembly

Trade‐offs maintain diversity and structure communities along environmental gradients. Theory indicates that if covariance among functional traits sets a limit on the number of viable trait combinations in a given environment, then communities with strong multidimensional trait constraints should exhibit low species diversity. We tested this prediction in winter annual plant assemblages along an aridity gradient using multilevel structural equation modelling. Univariate and multivariate functional diversity measures were poorly explained by aridity, and were surprisingly poor predictors of community richness. By contrast, the covariance between maximum height and seed mass strengthened along the aridity gradient, and was strongly associated with richness declines. Community richness had a positive effect on local neighbourhood richness, indicating that climate effects on trait covariance indirectly influence diversity at local scales. We present clear empirical evidence that declines in species richness along gradients of environmental stress can be due to increasing constraints on multidimensional phenotypes.     

Responses of plant functional types to an environmental gradient on the Northeast China Transect

The hypothesis that some plant traits such as life form are robust surrogates for plant functional type (PFT) has provoked an ongoing debate. Based on a dataset from the Northeast China Transect (NECT), we attempted to test the hypothesis by comparing an objective PFT identification framework in which large datasets of plant traits were considered with two subjective PFT frameworks in which only a few plant traits were involved. Additionally, we addressed the relations between the relative abundance of PFTs and the environmental gradient represented by actual evapotranspiration (AET) along the NECT. We also discuss the changes in ecosystem functioning associated with the PFT turnover along the environmental gradient. Based on an objective PFT classification, eight PFTs were identified: deciduous trees, shrubs, perennial forbs with lower net photosynthesis, perennial forbs with higher net photosynthesis, perennial bulb-grasses, perennial tiller-grasses, annual C₄ herbs and evergreen trees. Our results indicated that some plant traits, such as life form and photosynthesis pathway, are robust surrogates for PFTs, implying that subjective approaches to PFT classification are useful. Nonetheless, caution should be used during the classification of PFTs. The framework adopted for PFT classification should depend on the specific scientific issues being dealt with. It is therefore meaningless to pursue a general framework for the identification of PFTs even within given plant communities. On the other hand, our quantitative classification of PFTs confirmed recurrent patterns with respect to PFT turnover along an environmental gradient. Furthermore, with the turnover in PFT along the NECT from the west to the east, ecosystem properties such as productivity and carbon storage are predicted to decrease, while photosynthesis is predicted to increase, suggesting that PFT turnover would inevitably lead to changes in ecosystem functioning.     

Functional traits as indicators of fodder provision over a short time scale in species-rich grasslands

Background and Aims: Fodder provision in species-rich grasslands, i.e. herbage growth,proportion of leaf, and leaf and stem digestibility, is difficultto predict for short periods of time, such as between two defoliationsor less. The value of two methods based on plant traits forevaluating these agronomic properties was examined. Methods: One method is based on plant trait measurements on the plantcommunity (leaf dry matter content, plant height, floweringdate); the other is on vegetation composition expressed as plantfunctional types (acquisitive versus conservative PFTs) establishedby measuring leaf dry matter content on pure grass stands. Theexperiment consisted of 18 fields with three different defoliationregimes (combinations of cutting and grazing) and two levelsof fertilization. To establish a growth curve over the firstgrowth cycle, herbage was sampled about 10 times in spring. Key Results: Coefficients of correlation between agronomic properties ofthe vegetation and its functional composition were higher whenthe latter was assessed through PFT and an indicator of theplant nutrient status (Ni) instead of measured plant traits.The date at which the ceiling yield occurred for the standingherbage mass or only the leaf component, which varied by upto 500 degree-days between treatments, and the leaf proportion,depended entirely on the PFT, and largely so for the leaf digestibility.The standing herbage mass at the time of ceiling yield dependedonly on Ni, or mainly so in the case of the daily herbage growthrate. Similar plant digestibility between plant communitieswas found at flowering time, although there were big differencesin PFT composition. The shape of the growth curve was flatterwhen there was great functional diversity in the plant community. Conclusions: The PFT composition and the Ni were more reliable than the plantfunctional traits measured in the field for evaluating herbagegrowth pattern and digestibility in spring.     

Diversity patterns of plant functional types in relation to fire regime and previous land use in Mediterranean woodlands

Abstract. We explore patterns of diversity of plant functional types (PFTs) in Mediterranean communities subjected to landscape‐scale fire disturbances in a mosaic of uncultivated and old fields stands. We use regenerative and growth form attributes to establish two sets of PFTs of perennial species living in shrublands and pine forests of NE Spain. We test the following hypotheses: 1. Fire frequency decreases regenerative PFTs diversity by negatively selecting attributes with low regenerative efficiency. 2. Fire history has more influence on regenerative than on growth form PFTs. 3. The lowest diversity of growth form PFTs will be in old fields without recent fires. We surveyed stands of different combinations of fire and land use histories. Fire history included areas without fires in the last twenty years (unburned), sites burned in 1982 (1‐burned), and sites burned in 1982 and 1994 (2‐burned). Land use histories considered terraced old fields, and uncultivated stands on stony soils. We analysed patterns of PFT abundance and diversity at the stand level, and across the landscape (among stands absolute deviations from sample medians of the relative cover of PFTs). At the stand level, fire had more influence on the diversity patterns of regenerative PFTs than on growth form PFTs. Fire decreased the diversity of regenerative PFTs, due to the elimination of the species without effective mechanisms to post‐fire regeneration. This effect was not observed across the landscape, but seeders showed more variation in stands with longer history without fire. Land use contributed to explain the diversity patterns of growth form PFT (i.e. the number of growth form PFTs was lower in uncultivated, unburned sites), but it did not influence regenerative PFTs diversity. Patterns of PFTs diversity reflect the response to ecological processes operating at the landscape level. Overall, regenerative and growth form PFTs appear to be more sensitive to the fire history than to the past land use.     

Photosynthetic and morphological functional types from different steppe communities in Inner Mongolia,North China

Morphological functional types and photosynthetic pathway types were identified for the forage species from steppe communities in Inner Mongolia, China, using the data of both field survey and published papers. Seven typical steppe communities were selected to investigate the morphological functional type and photosynthetic pathway type compositions and plant functional type (PFT) diversity in steppe communities at regional scale. Morphological functional types, based on plant height and leaf type combined with life span, were optimal for comparing the community differences in the region, while photosynthetic pathway types were fairly coarse for such studies. Of the seven morphological functional types in the steppe communities, perennial forbs (PEF) were the dominant type, and 60 % of species belonged to this type. Each of the high perennial grass (HPG), short perennial grass (SPG), and annual grass (ANG) types represented less than 10 % of the total, even though the grass species were dominant in the seven steppe communities. The differences of PFTs between the steppe communities were remarkable, and the PFT richness and diversity increased from the communities with moist conditions to the ones with dry environments.     

Functional Trait Strategies of Trees in Dry and Wet Tropical Forests Are Similar but Differ in Their Consequences for Succession

Global plant trait studies have revealed fundamental trade-offs in plant resource economics. We evaluated such trait trade-offs during secondary succession in two species-rich tropical ecosystems that contrast in precipitation: dry deciduous and wet evergreen forests of Mexico. Species turnover with succession in dry forest largely relates to increasing water availability and in wet forest to decreasing light availability. We hypothesized that while functional trait trade-offs are similar in the two forest systems, the successful plant strategies in these communities will be different, as contrasting filters affect species turnover. Research was carried out in 15 dry secondary forest sites (5-63 years after abandonment) and in 17 wet secondary forest sites (<1-25 years after abandonment). We used 11 functional traits measured on 132 species to make species-trait PCA biplots for dry and wet forest and compare trait trade-offs. We evaluated whether multivariate plant strategies changed during succession, by calculating a ‘Community-Weighted Mean’ plant strategy, based on species scores on the first two PCA-axes. Trait spectra reflected two main trade-off axes that were similar for dry and wet forest species: acquisitive versus conservative species, and drought avoiding species versus evergreen species with large animal-dispersed seeds. These trait associations were consistent when accounting for evolutionary history. Successional changes in the most successful plant strategies reflected different functional trait spectra depending on the forest type. In dry forest the community changed from having drought avoiding strategies early in succession to increased abundance of evergreen strategies with larger seeds late in succession. In wet forest the community changed from species having mainly acquisitive strategies to those with more conservative strategies during succession. These strategy changes were explained by increasing water availability during dry forest succession and increasing light scarcity during wet forest succession. Although similar trait spectra were observed among dry and wet secondary forest species, the consequences for succession were different resulting from contrasting environmental filters.     

Functional Traits in a Species-Rich Grassland and a Short-Term Change in Management: Is There a Competition-Colonization Trade-Off?

The species richness of grasslands generally cannot be fully restored after changes in management. Some species with small statures and basal leaf rosettes can be lost forever. The same species, however, seem to possess the traits necessary for successful re-colonization – they produce small, easily dispersable seeds, numerous seedlings and have lasting seed banks. We tested the hypothesis that plants in species-rich grasslands can be characterized by a negative correlation between their competitive ability and potential for generative regeneration, i.e. by a competition-colonization trade-off. An analysis of the traits of 95 grassland species supported this hypothesis. We then conducted a manipulative experiment in three different meadow communities in the Bílé Karpaty Mts. The experiment involved characterizing species traits during periods of different grassland management regimes in the years 1997–2000 and comparing these with the original management regime, which was restored between 2000 and 2003. We found out that the hypothesis only holds true for the pooled dataset for all three communities. When the individual meadow communities were analyzed separately, plant traits other than those responsible for the competition-colonization trade-off appear to be characteristic of responsive species, e.g. shoot lifespan or phenology. Our results imply that despite the general trade-offs found in large comparative studies, the plant response in a specific community is constrained by the local species pool.     

Regeneration of different plant functional types in a Masson pine forest following pine wilt disease

Pine wilt disease is a severe threat to the native pine forests in East Asia. Understanding the natural regeneration of the forests disturbed by pine wilt disease is thus critical for the conservation of biodiversity in this realm. We studied the dynamics of composition and structure within different plant functional types (PFTs) in Masson pine forests affected by pine wilt disease (PWD). Based on plant traits, all species were assigned to four PFTs: evergreen woody species (PFT1), deciduous woody species (PFT2), herbs (PFT3), and ferns (PFT4). We analyzed the changes in these PFTs during the initial disturbance period and during post-disturbance regeneration. The species richness, abundance and basal area, as well as life-stage structure of the PFTs changed differently after pine wilt disease. The direction of plant community regeneration depended on the differential response of the PFTs. PFT1, which has a higher tolerance to disturbances, became dominant during the post-disturbance regeneration, and a young evergreen-broad-leaved forest developed quickly after PWD. Results also indicated that the impacts of PWD were dampened by the feedbacks between PFTs and the microclimate, in which PFT4 played an important ecological role. In conclusion, we propose management at the functional type level instead of at the population level as a promising approach in ecological restoration and biodiversity conservation.     

Photosynthetic and morphological functional types from different steppe communities in Inner Mongolia, North China

Morphological functional types and photosynthetic pathway types were identified for the forage species from steppe communities in Inner Mongolia, China, using the data of both field survey and published papers. Seven typical steppe communities were selected to investigate the morphological functional type and photosynthetic pathway type compositions and plant functional type (PFT) diversity in steppe communities at regional scale. Morphological functional types, based on plant height and leaf type combined with life span, were optimal for comparing the community differences in the region, while photosynthetic pathway types were fairly coarse for such studies. Of the seven morphological functional types in the steppe communities, perennial forbs (PEF) were the dominant type, and 60 % of species belonged to this type. Each of the high perennial grass (HPG), short perennial grass (SPG), and annual grass (ANG) types represented less than 10 % of the total, even though the grass species were dominant in the seven steppe communities. The differences of PFTs between the steppe communities were remarkable, and the PFT richness and diversity increased from the communities with moist conditions to the ones with dry environments.This revised version was published online in March 2005 with corrections to the page numbers.     

Clumped versus scattered: how does the spatial correlation of disturbance events affect biodiversity?

In this study, we systematically explore the effects of rate and spatial correlation (level of clumping) of disturbance events on a community of sessile species differing in their life history traits. A spatially explicit individual-based model shows that long-term coexistence is very sensitive to spatial correlation when the trade-off in life history traits includes differences in dispersal distances. Highest biodiversity emerges at highly correlated disturbances of intermediate rates. Diversity peaks shift to larger rates when clumping decreases. Scattered disturbances lead to competitive exclusion. Interestingly, we observed additional peaks in the diversity–disturbance curves at certain levels of clumping. Thus, subject to the differences in life history traits, particular combinations of disturbance rate and spatial correlation may enable subsets of species to coexist, which opens new possibilities for explaining diversity. Our results suggest that observation of high biodiversity under spatially correlated disturbances points to a competition–colonisation trade-off, which includes dispersal distances.     

Elevational trends of biodiversity and plant traits do not converge—a test in the Helan Range, NW China

When elevational gradients include combinations of different climatic gradients, such as a decline in temperature combined with an increase in moisture, vegetation and plant trait responses are difficult to explain. Here, we used plant species richness and morphometric traits data across steep elevational gradients in the Helan Range of Northwestern China in an attempt to separate general trends (temperature related) from regional peculiarity (moisture related). Based on the floristic data of the Helan Range as well as plot-based data, both drought (at low elevation) and low temperature (high elevation) are associated with low species richness presumably also explaining the peak in diversity at mid-elevation, where climatic conditions are moderate. However, this mid-elevation peak in diversity is not mirrored in trends of plant traits such as leaf size and inflorescence size, which show either unidirectional trends or no change with elevation (with impacts of drought and low temperature perhaps gradually replacing each other). Our analysis illustrated the taxonomic and plant functional type (PFT)-related biases in functional trait studies and showed that consistent patterns only emerge after careful data stratification, with taxonomy (family level) holding more promises than PFTs. Inflorescence size increased with elevation in major insect-pollinated families, a trend not seen in wind-pollinated (graminoid) families. However, the reproductive effort expressed as inflorescence/leaf length ratio increases with elevation in the majority of plant families, irrespective of their pollination system. The fact that these biometric responses to elevation do not correlate with responses in species richness (which peaks at mid-elevation) may reflect contrasting drivers of trait selection and biodiversity. Based on our plot-based data, this analysis also confirmed the usefulness of floristic archive data for testing ecological theory related to elevational gradients.     

A trait-based approach for modelling microbial litter decomposition

Trait-based models are an emerging tool in ecology with the potential to link community dynamics, environmental responses and ecosystem processes. These models represent complex communities by defining taxa with trait combinations derived from prior distributions that may be constrained by trade-offs. Herein I develop a model that links microbial community composition with physiological and enzymatic traits to predict litter decomposition rates. This approach allows for trade-offs among traits that represent alternative microbial strategies for resource acquisition. The model predicts that optimal strategies depend on the level of enzyme production in the whole community, which determines resource availability and decomposition rates. There is also evidence for facilitation and competition among microbial taxa that co-occur on decomposing litter. These interactions vary with community investment in extracellular enzyme production and the magnitude of trade-offs affecting enzyme biochemical traits. The model accounted for 69% of the variation in decomposition rates of 15 Hawaiian litter types and up to 26% of the variation in enzyme activities. By explicitly representing diversity, trait-based models can predict ecosystem processes based on functional trait distributions in a community. The model developed herein illustrates that traits influencing microbial enzyme production are some of the key controls on litter decomposition rates.     

Functional flower traits and their diversity drive pollinator visitation

Recent studies have shown that the diversity of flowering plants can enhance pollinator richness and visitation frequency and thereby increase the resilience of pollination. It is assumed that flower traits explain these effects, but it is still unclear which flower traits are responsible, and knowing that, if pollinator richness and visitation frequency are more driven by mass‐ratio effects (mean trait values) or by trait diversity. Here, we analyse a three‐year data set of pollinator observations collected in a European grassland plant diversity experiment (The Jena experiment). The data entail comprehensive flower trait measurements, including reward traits (nectar and pollen amount), morphological traits (height, symmetry, area, colour spectra) and chemical traits (nectar‐amino acid and nectar‐sugar concentration). We test if pollinator species richness and visitation frequency of flower communities depend on overall functional diversity combining all flower traits within a community, single trait diversities (within trait variation) and community‐weighted means of the single traits, using Bayesian inference. Overall functional diversity did not affect pollinator species richness, but reduced visitation frequency. When looking at individual flower traits separately, we found that single trait diversity of flower reflectance and flower morphology were important predictors of pollinator visitation frequency. Moreover, independent of total flower abundance, community‐weighted means of flower height, area, reflectance, nectar‐sugar concentration and nectar‐amino acid concentration strongly affected both pollinator species richness and visitation frequency. Our results, challenge the idea that functional diversity always positively affects ecosystem functions. Nonetheless, we demonstrate that both single trait diversity and mass‐ratio effects of flower traits play an important role for diverse and frequent flower visits, which underlines the functionality of flower traits for pollination services.     

Four decades of functional community change reveals gradual trends and low interlinkage across trophic groups in a large marine ecosystem

The rate at which biological diversity is altered on both land and in the sea, makes temporal community development a critical and fundamental part of understanding global change. With advancements in trait‐based approaches, the focus on the impact of temporal change has shifted towards its potential effects on the functioning of the ecosystems. Our mechanistic understanding of and ability to predict community change is still impeded by the lack of knowledge in long‐term functional dynamics that span several trophic levels. To address this, we assessed species richness and multiple dimensions of functional diversity and dynamics of two interacting key organism groups in the marine food web: fish and zoobenthos. We utilized unique time series‐data spanning four decades, from three environmentally distinct coastal areas in the Baltic Sea, and assembled trait information on six traits per organism group covering aspects of feeding, living habit, reproduction and life history. We identified gradual long‐term trends, rather than abrupt changes in functional diversity (trait richness, evenness, dispersion) trait turnover, and overall multi‐trait community composition. The linkage between fish and zoobenthic functional community change, in terms of correlation in long‐term trends, was weak, with timing of changes being area and trophic group specific. Developments of fish and zoobenthos traits, particularly size (increase in small size for both groups) and feeding habits (e.g. increase in generalist feeding for fish and scavenging or predation for zoobenthos), suggest changes in trophic pathways. We summarize our findings by highlighting three key aspects for understanding functional change across trophic groups: (a) decoupling of species from trait richness, (b) decoupling of richness from density and (c) determining of turnover and multi‐trait dynamics. We therefore argue for quantifying change in multiple functional measures to help assessments of biodiversity change move beyond taxonomy and single trophic groups.     

The relative importance of local, regional and historical factors determining the distribution of plants in fragmented riverine forests: an emergent group approach

Aim To investigate how local, regional and historical factors shape the herbaceous plant communities in fragmented riverine forests, and how the community composition and species richness of these fragments is related to the interplay between the environmental factors and specific plant life‐trait combinations. Location Riverine forest fragments in the Grand‐duché de Luxembourg. Methods Forest fragments were surveyed for their abundance in herbaceous plant species. All plant species where clustered into Emergent Groups (EG) by means of a formal classification based on 14 life‐history traits. Within each EG, the local, regional and historical factors were related to the community composition using partial Canonical Correspondence Analyses (pCCA) and to the species richness using Generalized Linear Models (GLMs). The EG colonization ability was characterized by means of logistic regressions. Results We defined and characterized seven EGs, among which three consisted of forest specialist species (barochorous perennials, short geophytes and zoochorous perennials), which exhibited specific life‐trait combinations: large and short‐lived seeds and/or vernal phenology. Differences in EG composition between forest fragments were mainly explained by local environmental factors such as soil productivity and pH. The richness of barochorous perennials and short geophytes was well predicted by the historical and regional factors. The colonization ability appeared very low for barochorous perennials and short geophytes. Main conclusions Local environmental conditions appear to drive the differentiation of the riverine forest plant communities owing to the specific habitat requirements of many forest species. Spatial and temporal forest discontinuities affect the richness of forest specialist species, due to dispersal and/or recruitment limitations. The emergent group approach enhances the understanding of the relative influence of local, regional and historical factors by distinguishing between forest specialists from generalists or ‘matrix’ species, which have a masking effect.     

南亚热带森林常见种和偶见种的功能多样性特征

群落特征可能取决于物种在功能实体功能特征的独特组合（FE）中的分布。了解这些分布是保护生物多样性和维持生态系统功能的基础。鼎湖山南亚热带常绿阔叶林群落物种丰富,根据物种的多度来区分常见种（每hm²超过一个个体）和偶见种（每hm²少于一个个体）。基于功能实体,分别计算功能实体数量、功能冗余度、功能脆弱度和功能过度冗余度来描述每个样方中常见种和偶见种的功能多样性。采用Wilcoxon检验检测常见种和偶见种功能性状的差异,通过线性回归分析常见种和偶见种的功能多样性与物种数量的关系,采用冗余分析（RDA）影响常见种和偶见种的功能多样性的环境因子。研究结果表明：（1）鼎湖山南亚热带常绿阔叶林群落中功能性状在常见种和偶见种间差异显著。（2）群落中数量较低的偶见种保持着较高比例的功能实体数量。功能实体数量、功能冗余度与常见种和偶见种数量显著正相关,功能过度冗余度与偶见种数量显著正相关,功能脆弱度与常见种和偶见种的物种数量呈显著负相关。（3）海拔是影响常见种功能多样性最关键的地形因子,凹凸度是影响偶见种功能多样性最关键的地形因子。土壤含水量是影响常见种功能多样性最关键的土壤因子,速效钾是影响偶见种功能多样性最关键的环境因子。研究结果表明,尽管物种丰富的热带森林具有较高的功能冗余,但其提供的保险效应不能抵消生态系统的功能脆弱性。减少具有独特功能实体和偶见种的损失是避免热带森林生态系统功能损失的有效和必要的方法。