Questions: The co-existence of high numbers of species has always fascinated ecologists, but what and where are the communities with the world records for plant species richness? The species—area relationship is among the best-known patterns in community ecology, but does it give a consistent global pattern for the most saturated communities, the global maxima? Location: The world. Methods: We assembled the maximum values recorded for vascular plant species richness for contiguous areas from 1 mm2 up to 1 ha. We applied the power function to relate maximal richness to area and to make extrapolations to the whole Earth. Results: Only two community types contain global plant species maxima. The maxima at smaller spatial grain were from oligo- to meso-trophic, managed, semi-natural, temperate grasslands (e.g. 89 species on 1 m 2 ), those at larger grains were from tropical rain forests (e.g. 942 species on 1 ha). The maximum richness values closely followed a power function with z = 0.250: close to Preston's 'canonical' value of 0.262. There was no discernable difference between maxima using rooted presence (i.e. including only plants rooted in the plot) vs shoot presence (i.e. including any plant with physical cover over the plot). However, shoot presence values must logically be greater, with the curves flattening out at very small grain, and there is evidence of this from point quadrats. Extrapolating the curve to the terrestrial surface of the Earth gave a prediction of 219 204 vascular plant species, surprisingly close to a recent estimate of 275 000 actual species. Conclusions: Very high richness at any spatial grain is found only in two particular habitat/community types. Nevertheless, these high richness values form a very strong, consistent pattern, not greatly affected by the method of sampling, and this pattern extrapolates amazingly well. The records challenge ecologists to consider mechanisms of species co-existence, answers to the 'Paradox of the Plankton'.
Question: Are plant traits more closely correlated with mean annual temperature, orwithmean annual precipitation? Location: Global. Methods: We quantified the strength of the relationships between temperature and precipitation and 21 plant traits from 447,961 species-site combinations worldwide. We used meta-analysis to provide an overall answer to our question. Results: Mean annual temperature was significantly more strongly correlated with plant traits than was mean annual precipitation. Conclusions: Our study provides support for some of the assumptions of classical vegetation theory, and points to many interesting directions for future research. The relatively low R
Question: Which functional diversity indices have the power to reveal changes in community assembly processes along abiotic stress gradients? Is their power affected by stochastic processes and variations in species richness along stress gradients? Methods: We used a simple community assembly model to explore the power of functional diversity indices across a wide range of ecological contexts. The model assumes that with declining stress the influence of niche complementarity on species fitness increases while that of environmental filtering decreases. We separately incorporated two trait-independent stochastic processes — mass and priority effects — in simulating species occurrences and abundances along a hypothetical stress gradient. We ran simulations where species richness was constant along the gradient, or increased, decreased or varied randomly with declining stress. We compared observed values for two indices of functional richness — total functional dendrogram length (FD) and convex hull volume (FRic) — with a matrix-swap null model (yielding indices SESFD and SESFRic) to remove any trivial effects of species richness. We also compared two indices that measure both functional richness and functional divergence — Rao quadratic entropy (Rao) and functional dispersion (FDis) — with a null model that randomizes abundances across species but within communities. This converts them to pure measures of functional divergence (SESRao and SESFDis). Results: When mass effects operated, only SESRao and SESFDis gave reasonable power, irrespective of how species richness varied along the stress gradient. FD, FRic, Rao and FDis had low power when species richness was constant, and variation in species richness greatly influenced their power. SESFRic and SESFD were unaffected by variation in species richness. When priority effects operated, FRic, SESFRic, Rao and FDis had good power and were unaffected by variation in species richness. Variation in species richness greatly affected FD and SESFD. SESRao and SESFDis had low power in the priority effects model but were unaffected by variation in species richness. Conclusions: Our results demonstrate that a reliable test for changes in assembly processes along stress gradients requires functional diversity indices measuring either functional richness or functional divergence. We recommend using SESFRic as a measure of functional richness and either SESRao or SESFDis (which are very closely related mathematically) as a measure of functional divergence. Used together, these indices of functional richness and functional divergence provide good power to test for increasing niche complementarity with declining stress across a broad range of ecological contexts.
Questions: What historical natural disturbances have shaped the structure and development of an old-growth, sub-alpine Picea abies forest? Are large-scale, high-severity disturbances (similar to the recent windthrow and bark beetle outbreaks in the region) within the historical range of variability for this forest ecosystem? Can past disturbances explain the previously described gradient in stand structure that had been attributed to an elevation gradient? Location: Šumava National Park (the Bohemian Forest) of the southwest Czech Republic. Methods: We reconstructed the site's disturbance history using dendroecological methods in a 20-ha study plot, established to span an elevation gradient. Growth patterns of 400 increment cores were screened for: (1 ) abrupt increases in radial growth indicating mortality of a former canopy tree and (2) rapid early growth rates indicating establishment in a former canopy gap. Results: Spatial and temporal patterns of canopy accession varied markedly over the 20-ha study area, resulting in disturbance pulses that corresponded to an elevation gradient. On the lower slope of the plot, the majority of the trees reached the canopy during two pulses (1770-1800 and 1820-1840), while most trees on the upper slope accessed the canopy in one pulse (1840-1860). Historically documented windstorms roughly coincide with peaks in our disturbance reconstruction. Conclusions: Our study provides strong evidence that these forests were historically shaped by infrequent, moderate- to high-severity natural disturbances. Our methods, however, could not definitively identify the agent(s) of these disturbances. Nevertheless, the recent mid-1990s windstorm and the ensuing spruce bark beetle outbreak may provide an analogue for past disturbance, as the duration and severity of these events could easily explain past patterns of growth response and recruitment in our results. Thus, it seems reasonable to assume the interaction of windstorms and bark beetles seen in the contemporary landscape has occurred historically. Finally, our results suggest that the previously documented elevation gradient in forest structure may not be related to elevation per se (lower temperatures and shorter growing season) but rather to changes in disturbance severity mediated by elevation.
Question: How many vegetation plot observations (relevés) are available in electronic databases, how are they geographically distributed, what are their properties and how might they be discovered and located for research and application? Location: Global. Methods: We compiled the Global Index of Vegetation-Plot Databases (GIVD; http://www.givd.info), an Internet resource aimed at registering metadata on existing vegetation databases. For inclusion, databases need to (i) contain temporally and spatially explicit species co-occurrence data and (ii) be accessible to the scientific public. This paper summarizes structure and data quality of databases registered in GIVD as of 30 December 2010. Results: On the given date, 132 databases containing more than 2.4 million non-overlapping plots had been registered in GIVD. The majority of these data were in European databases (83 databases, 1.6 million plots), whereas other continents were represented by substantially less (North America 15, Asia 13, Africa nine, South America seven, Australasia two, multi-continental three). The oldest plot observation was 1864, but most plots were recorded after 1970. Most plots reported vegetation on areas of 1 to 1000 m2; some also stored time-series and nested-plot data. Apart from geographic reference (required for inclusion), most frequent information was on altitude (71%), slope aspect and inclination (58%) and land use (38%), but rarely soil properties ( < 7%). Conclusions: The vegetation plot data in GIVD constitute a major resource for biodiversity research, both through the large number of species occurrence records and storage of species co-occurrence information at a small scale, combined with structural and plot-based environmental data. We identify shortcomings in available data that need to be addressed through sampling under-represented geographic regions, providing better incentives for data collection and sharing, developing user-friendly database exchange standards, as well as tools to analyse and remove confounding effects of sampling biases. The increased availability of data sets conferred by registration in GIVD offers significant opportunities for large-scale studies in community ecology, macroecology and global change research.
A novel conceptual framework is presented that proposes to apply trait-based approaches to predicting the impact of environmental change on ecosystem service delivery by multi-trophic systems. Development of the framework was based on an extension of the response—effect trait approach to capture functional relationships that drive trophic interactions. The framework was populated with worked examples to demonstrate its flexibility and value for linking disparate data sources, identifying knowledge gaps and generating hypotheses for quantitative models.
Question: Mean Ellenberg indicator values (EIVs) inherit information about compositional similarity, because during their calculation species abundances (or presence—absences) are used as weights. Can this similarity issue actually be demonstrated, does it bias results of vegetation analyses correlating mean EIVs with other aspects of species composition and how often are biased studies published? Methods: In order to separate information on compositional similarity possibly present in mean EIVs, a new variable was introduced, calculated as a weighted average of randomized species EIVs. The performance of these mean randomized EIVs was compared with that of the mean real EIVs on the one hand and random values (randomized mean EIVs) on the other. To demonstrate the similarity issue, differences between samples were correlated with dissimilarity matrices based on various indices. Next, the three mean EIV variables were tested in canonical correspondence analysis (CCA), detrended correspondence analysis (DCA), analysis of variance (ANOVA) between vegetation clusters, and in regression on species richness. Subsequently, a modified permutation test of significance was proposed, taking the similarity issue into account. In addition, an inventory was made of studies published in the Journal of Vegetation Science and Applied Vegetation Science between 2000 and 2010 likely reporting biased results due to the similarity issue. Results: Using mean randomized EIVs, it is shown that compositional similarity is inherited into mean EIVs and most resembles the inter-sample distances in correspondence analysis, which itself is based on iterative weighted averaging. The use of mean EIVs produced biased results in all four analysis types examined: unrealistic (too high) explained variances in CCA, too many significant correlations with ordination axes in DCA, too many significant differences between cluster analysis groups and too high coefficients of determination in regressions on species richness. Modified permutation tests provided ecologically better interpretable results. From 95 studies using Ellenberg indicator values, 36 reported potentially biased results. Conclusions: No statistical inferences should be made in analyses relating mean EIVs with other variables derived from the species composition as this can produce highly biased results, leading to misinterpretation. Alternatively, a modified permutation test using mean randomized EIVs can sometimes be used.
Question: How do multivariate methods perform in relating species- and community-level trait responses to the environment? Location: (1) Field data from grazed semi-natural grasslands, NE Germany; (2) artificial data. Methods: Research questions associated with trait—environment relationships were briefly reviewed and seven available methods evaluated. The main distinction between research questions is whether trait—environment relationships should be addressed at community or species level. A redundancy analysis (RDA) of mean trait values of species in a plot weighted by their abundances (CWM-RDA) is exclusively suitable for the community level. The other six methods address the species level. A double inertia analysis of two arrays (RLQ) and double canonical correspondence analysis (double CCA) use combinations of ordinations to simultaneously analyse species and trait responses to the environment. A combination of the outlying mean index with generalized additive models (OMI-GAM) predicts the response of species to environmental variables on trait gradients. RDA-RegTree first analyses species responses to the environment with RDA and then uses a regression tree to classify trait expressions according to scores of species responses on the ordination axes. Cluster regression uses cluster analyses and logistic regression to search for trait combinations with the best response to the environmental variables. This method models the distribution of functional groups on environmental gradients. All methods and data are available as R scripts. Results: All methods consistently revealed the main trait responses to environment in the field data set, namely that life history was associated with available phosphorus while grazing intensity was related to leaf C:N ratio and canopy height. At community level, CWM-RDA gave a good overview of trait— environment relationships, as also provided by the species-based methods RLQ and double CCA. OMI-GAM revealed non-linear relationships in the field data set. Field and artificial data gave that the number and stability of functional groups produced by Cluster regression and RDA-RegTree varied more strongly than RLQ, double CCA and OMI-GAM. Conclusions: Each method addresses particular ecological concepts and research questions. If a user asks for the response of average trait expressions of communities to environmental gradients, CWM-RDA may be the first choice. However, species-based methods should be applied to address questions regarding co-existence of different life histories or to assess how groups of species respond to environmental changes. The artificial data set revealed that the methods differed in sensitivity to gradient lengths and random data.
Background: In forests subject to stand-replacing disturbances, conventional models of succession typically overlook early-seral stages as a simple re-organization/establishment period. These models treat structural development in essentially 'relay floristic' terms, with structural complexity (three-dimensional heterogeneity) developing primarily in old-growth stages, only after a closedcanopy 'self-thinning' phase and subsequent canopy gap formation. However, is it possible that early-successional forests can sometimes exhibit spatial complexity similar to that in old-growth forests — i.e. akin to an 'initial floristic' model of structural development? Hypothesis: Based on empirical observations, we present a hypothesis regarding an important alternative pathway in which protracted or sparse forest establishment and interspecific competition thin out tree densities early on — thereby precluding overstorey canopy closure or a traditionally defined self-thinning phase. Although historically viewed as an impediment to stand development, we suggest this process may actually advance certain forms of structural complexity. These young stands can exhibit qualities typically attributed only to old forests, including: (1) canopy gaps associated with clumped and widely spaced tree stems; (2) vertically heterogeneous canopies including under- and midstories, albeit lower stature; (3) co-existence of shade-tolerant and intolerant species; and (4) abundant dead wood. Moreover, some of these qualities may persist through succession, meaning that a significant portion of eventual old-growth spatial pattern may already be determined in this early stage. Implications: The relative frequency of this open-canopy pathway, and the degree to which precocious complexity supports functional complexity analogous to that of old forests, are largely unknown due to the paucity of naturally regenerating forests in many regions. Nevertheless, recognition of this potential is important for the understanding and management of early-successional forests.
Questions: Did the forest area in the Swiss Alps increase between 1985 and 1997? Does the forest expansion near the tree line represent an invasion into abandoned grasslands (ingrowth) or a true upward shift of the local tree line? What land cover / land use classes did primarily regenerate to forest, and what forest structural types did primarily regenerate? And, what are possible drivers of forest regeneration in the tree line ecotone, climate and/or land use change?Location: Swiss Alps.Methods: Forest expansion was quantified using data from the repeated Swiss land use statistics GEOSTAT. A moving window algorithm was developed to distinguish between forest ingrowth and upward shift. To test a possible climate change influence, the resulting upward shifts were compared to a potential regional tree line.Results: A significant increase of forest cover was found between 1650 m and 2450 m. Above 1650 m, 10% of the new forest areas were identified as true upward shifts whereas 90% represented ingrowth, and we identified both land use and climate change as likely drivers. Most upward shift activities were found to occur within a band of 300 m below the potential regional tree line, indicating land use as the most likely driver. Only 4% of the upward shifts were identified to rise above the potential regional tree line, thus indicating climate change.Conclusions: Land abandonment was the most dominant driver for the establishment of new forest areas, even at the tree line ecotone. However, a small fraction of upwards shift can be attributed to the recent climate warming, a fraction that is likely to increase further if climate continues to warm, and with a longer time-span between warming and measurement of forest cover.
Analyses of functional traits have become fundamental tools for understanding patterns and processes in plant community ecology. In this context, regenerative seed traits play an important, yet overlooked, role because they largely determine the ability of plants to disperse and re-establish. A survey of recent publications in community ecology suggests that seed germination traits in particular are neglected at the expense of other relevant but overused traits based only on seed morphology. As a response to this bias, we discuss the functional significance of seed germination traits in comparison with morphological and biophysical seed traits, and advocate their use in vegetation science. We also demonstrate how research in community assembly, climate change and restoration ecology can benefit from the inclusion of germination traits, encompassing functions that cannot be explained solely by adult plant traits. Seed germination experiments conducted in the laboratory or field to quantify these traits provide ecologically meaningful and relatively easy-to-obtain information about the functional properties of plant communities. We argue that bridging the gap between seed physiologists and community ecologists will improve the prediction of plant assemblages, and propose further perspectives for including seed traits into the research agenda of functional community ecologists.
In landscapes subject to intensive agriculture, both soil fertility and vegetation disturbance are capable of impacting strongly, evenly and simultaneously on the herbaceous plant cover and each tends to impose uniformity on the traits of constituent species. In more natural and ancient grasslands greater spatial and temporal variation in both productivity and disturbance occurs and both factors have been implicated in the maintenance of species-richness in herbaceous communities. However, empirical data suggest that disturbance is the more potent driver of trait differentiation and species co-existence at a local scale. This may arise from the great diversity in opportunities for establishment, growth or reproduction that arise when the intensity of competition is reduced by damage to the vegetation.In contrast to the diversifying effects of local disturbances, productivity-related plant traits (growth rate, leaf longevity, leaf chemistry, leaf toughness, decomposition rate) appear to be less variable on a local scale. This difference in the effects of the productivity and disturbance filters arises from the relative constancy of productivity within the community and the diversity in agency and in spatial and temporal scales exhibited by disturbance events. Also, evolutionary responses to disturbances involve minor adaptive shifts in phenological and regenerative traits and are more likely to occur as micro-evolutionary steps than the shifts in linked traits in the core physiology associated with the capacity to exploit productive and unproductive habitats.During the assembly of a community and over its subsequent lifespan filters with diversifying and convergent effects may operate simultaneously on recruitment from the local species pool and impose contrasted effects on the similarity of the trait values exhibited by co-existing species. Moreover, as a consequence of the frequent association of productivity with the convergence filter, an additional difference is predicted in terms of the effects of the two filters on ecosystem functioning. Convergence in traits selected by the productivity filter will exert effects on both the plant community and the ecosystem while divergent effects of the disturbance filter will be restricted to the plant community.
A methodology for partitioning of biodiversity into α, β and γ components has long been debated, resulting in different mathematical frameworks. Recently, use of the Rao quadratic entropy index has been advocated since it allows comparison of various facets of diversity (e. g. taxonomic, phylogenetic and functional) within the same mathematical framework. However, if not well implemented, the Rao index can easily yield biologically meaningless results and lead into a mathematical labyrinth. As a practical guideline for ecologists, we present a critical synthesis of diverging implementations of the index in the recent literature and a new extension of the index for measuring β-diversity. First, we detail correct computation of the index that needs to be applied in order not to obtain negative β-diversity values, which are ecologically unacceptable, and elucidate the main approaches to calculate the Rao quadratic entropy at different spatial scales. Then, we emphasize that, similar to other entropy measures, the Rao index often produces lower-than-expected β-diversity values. To solve this, we extend a correction based on equivalent numbers, as proposed by Jost (2007), to the Rao index. We further show that this correction can be applied to additive partitioning of diversity and not only its multiplicative form. These developments around the Rao index open up an exciting avenue to develop an estimator of turnover diversity across different environmental and temporal scales, allowing meaningful comparisons of partitioning across species, phylogenetic and functional diversities within the same mathematical framework. We also propose a set of R functions, based on existing developments, which perform different key computations to apply this framework in biodiversity science.
Fig. The three data sets used in the present study. We tested if the magnitude of intraspecific variability of co‐occurring species was lower than interspecific variability for a given trait by combining different approaches commonly used in functional ecology ‐ i.e., experimental data, field observations and extraction of data from a global database. We found that interspecific variability was greater than intraspecific variability and species ranking was conserved across different datasets and spatial scales.
Co‐existence theories fail to adequately explain observed community patterns (diversity and composition) because they mainly address local extinctions. For a more complete understanding, the regional processes responsible for species formation and geographic dispersal should also be considered. The species pool concept holds that local variation in community patterns is dependent primarily on the availability of species, which is driven by historical diversification and dispersal at continental and landscape scales. However, empirical evidence of historical effects is limited. This slow progress can be attributed to methodological difficulties in determining the characteristics of historical species pools and how they contributed to diversity patterns in contemporary landscapes. A role of landscape‐scale dispersal limitation in determining local community patterns has been demonstrated by numerous seed addition experiments. However, disentangling general patterns of dispersal limitation in communities still requires attention. Distinguishing habitat‐specific species pools can help to meet both applied and theoretical challenges. In conservation biology, the use of absolute richness may be uninformative because the size of species pools varies between habitats. For characterizing the dynamic state of individual communities, biodiversity relative to species pools provides a balanced way of assessing communities in different habitats. Information about species pools may also be useful when studying community assembly rules, because it enables a possible mechanism of trait convergence (habitat filtering) to be explicitly assessed. Empirical study of the role of historic effects and dispersal on local community patterns has often been restricted due to methodological difficulties in determining habitat‐specific species pools. However, accumulating distributional, ecological and phylogenetic information, as well as use of appropriate model systems (e.g. archipelagos with known biogeographic histories) will allow the species pool concept to be applied effectively in future research. In order to explain observed community patterns, the regional processes responsible for species formation and geographic dispersal should also be considered. The species pool concept offers a frame for that. It holds that local variation in community patterns is dependent primarily on the availability of species, which is driven by historical diversification and dispersal at continental and landscape scales.
Question: A set of easily-measured ('soft') plant traits has been identified as potentially useful predictors of ecosystem functioning in previous studies. Here we aimed to discover whether the screening techniques remain operational in widely contrasted circumstances, to test for the existence of axes of variation in the particular sets of traits, and to test for their links with 'harder' traits of proven importance to ecosystem functioning. Location: central-western Argentina, central England, northern upland Iran, and north-eastern Spain. Recurrent patterns of ecological specialization: Through ordination of a matrix of 640 vascular plant taxa by 12 standardized traits, we detected similar patterns of specialization in the four floras. The first PCA axis was identified as an axis of resource capture, usage and release. PCA axis 2 appeared to be a size-related axis. Individual PCA for each country showed that the same traits remained valuable as predictors of resource capture and utilization in all of them, despite their major differences in climate, biogeography and land-use. The results were not significantly driven by particular taxa: the main traits determining PCA axis 1 were very similar in eudicotyledons and monocotyledons and Asteraceae, Fabaceae and Poaceae. Links between recurrent suites of 'soft' traits and 'hard' traits: The validity of PCA axis 1 as a key predictor of resource capture and utilization was tested by comparisons between this axis and values of more rigorously established predictors ('hard' traits) for the floras of Argentina and England. PCA axis 1 was correlated with variation in relative growth rate, leaf nitrogen content, and litter decomposition rate. It also coincided with palatability to model generalist herbivores. Therefore, location on PCA axis 1 can be linked to major ecosystem processes in those habitats where the plants are dominant. Conclusion: We confirm the existence at the global scale of a major axis of evolutionary specialization, previously recognised in several local floras. This axis reflects a fundamental trade-off between rapid acquisition of resources and conservation of resources within well-protected tissues. These major trends of specialization were maintained across different environmental situations (including differences in the proximate causes of low productivity, i.e. drought or mineral nutrient deficiency). The trends were also consistent across floras and major phylogenetic groups, and were linked with traits directly relevant to ecosystem processes.
How have the frequency and severity of natural disturbances in primary Picea abies forests varied during recent centuries? The disturbance regime is characterized by considerable spatial and temporal heterogeneity, which could be seen among plots within stands, among stands within landscapes. When the disturbance regime was evaluated at larger scales, the entire range of disturbance severity was revealed.
An enormous recent research effort focused on how plant biodiversity (notably species richness) influences ecosystem functioning, usually through experiments in which diversity is varied through random draws of species from a species pool. Such experiments are increasingly used to predict how species losses influence ecosystem functioning in real' ecosystems. However, this assumes that comparisons of experimental communities with low vs high species richness are analogous to comparisons of natural communities from which species either have or have not been lost. I explore the validity of this assumption, and highlight difficulties in using such experiments to draw conclusions about the ecosystem consequences of biodiversity loss in natural systems. Notably, these experiments do not mimic what happens in real ecosystems either when local extinctions occur or when species losses are offset by gains of new species. Despite limitations, this single experimental approach for studying how biodiversity loss affects ecosystems has often been advocated and implemented at the expense of other approaches; this limits understanding of how natural ecosystems respond to biodiversity loss. I conclude that a broader spectrum of approaches, and more explicit consideration of how species losses and gains operate in concert to influence ecosystems, will help progress this field.
Clonal growth of plants is mainly a result of the vegetative growth of organs hidden beneath the soil surface and producing potentially independent vegetative offspring. Clonal traits are difficult to measure due to inaccessibility of the space they inhabit and their morphological diversity. This causes great difficulties with descriptions, standardization of measurements across plant growth forms and, probably, a lack of appropriate questions that should be answered using them. The freely available CLO-PLA database (http://clopla.butbn.cas.cz/) can help to assess the roles of vegetative means of regeneration and spread in plant communities under the effect of various biotic and abiotic filters. It can serve as a source of reference on persistence traits of European temperate flora and, eventually, as a guide for trait sampling in other regions of the world.
Despite the importance of measuring tropical forest biomass, the accuracy of biomass estimates is poorly constrained due to fundamental weaknesses in the design and implementation of field studies. We identify these issues and propose a radical paradigm shift to advance tropical forest biomass research to a firmer theoretical and empirical basis.