Plasticity in plant functional traits is shaped by variability in neighbourhood species composition.

Plant functional traits can vary widely as a result of phenotypic plasticity to abiotic conditions. Trait variation may also reflect responses to the identity of neighbours, although not all species are equally responsive to their biotic surroundings. We hypothesized that responses to neighbours are shaped by spatial community patterns and resulting variability in neighbour composition. More precisely, we tested the theoretical prediction that plasticity is most likely to evolve if alternative environments (in this case, different neighbour species) are common and encountered at similar frequencies. We estimated the frequencies of encountering different neighbour species in the field for 27 grassland species and measured the aboveground morphological responses of each species to conspecific vs heterospecific neighbours in a common garden. Responses to neighbour identity were dependent on how frequently the experimental neighbours were encountered by the focal species in their home community, with the greatest plasticity observed in species that encountered both neighbours (conspecific and heterospecific) with high and even frequency. Biotic interactions with neighbouring species can impose selection on plasticity in functional traits, which may feed back through trait divergence and niche differentiation to influence species coexistence and community structure.

Is the positive relationship between species richness and shoot morphological plasticity mediated by ramet density or is there a direct link?

Little is known about the consequences of phenotypic plasticity in co-existing species for plant community structure. However, it has been proposed that the potential of plants to exhibit plastic responses to light availability could be a key factor determining the capability of individuals to co-exist at small scales. Our previous research demonstrated that morphological plasticity to light was positively related to small-scale species richness in a temperate grassland. However, it remained unclear whether this relationship was solely due to a higher shoot density in plastic assemblages, or whether diversity was directly related to the morphological plasticity of the co-inhabitants. We used two data sets to clarify this relationship: experimentally acquired estimates of plasticity to light availability for 45 herbaceous plant species, and species richness and ramet density data from a 2-year permanent plot study in a semi-natural calcareous grassland. There was little ramet mortality observed in the permanent plot study indicating that the link between plasticity and richness does not operate through reduced mortality in more morphologically plastic assemblages. The local density of ramets explained most of variation in small-scale richness, but there was also a significant direct density-independent effect of mean shoot plasticity on richness, showing that plasticity to light directly enhances the small-scale co-existence of species.

Functional species pool framework to test for biotic effects on community assembly.

Functional trait differences among species are increasingly used to infer the effects of biotic and abiotic processes on species coexistence. Commonly, the trait diversity observed within communities is compared to patterns simulated in randomly generated communities based on sampling within a region. The resulting patterns of trait convergence and divergence are assumed to reveal abiotic and biotic processes, respectively. However, biotic processes such as competition can produce both trait divergence and convergence, through either excluding similar species (niche differences, divergence) or excluding dissimilar species (weaker competitor exclusion, convergence). Hence, separating biotic and abiotic processes that can produce identical patterns of trait diversity, or even patterns that neutralize each other, is not feasible with previous methods. We propose an operational framework in which the functional trait dissimilarity within communities (FDcomm) is compared to the corresponding trait dissimilarity expected from the species pool (i.e., functional species pool diversity, FDpool). FDpool includes the set of potential species for a site delimited by the operating environmental and dispersal limitation filters. By applying these filters, the resulting pattern of trait diversity is consistent with biotic processes, i.e., trait divergence (FDcomm > FDpool) indicates niche differentiation, while trait convergence (FDcomm < FDpool) indicates weaker competitor exclusion. To illustrate this framework, with its potential application and constraints, we analyzed both simulated and field data. The functional species pool framework more consistently detected the simulated trait diversity patterns than previous approaches. In the field, using data from plant communities of typical Northern European habitats in Estonia, we found that both niche-based and weaker competitor exclusion influenced community assembly, depending on the traits and community considered. In both simulated and field data, we demonstrated that only by estimating the species pool of a site is it possible to differentiate the patterns of trait dissimilarity produced by operating biotic processes. The framework, which can be applied with both functional and phylogenetic diversity, enables a reinterpretation of community assembly processes. Solving the challenge of defining an appropriate reference species pool for a site can provide a better understanding of community assembly.

Understory plant diversity is related to higher variability of vegetative mobility of coexisting species.

Theoretical studies claim that if co-occurring species have very different mobilities this will result in greater small-scale species richness, but empirical evidence is still lacking. We measured horizontal vegetative mobility (VM) of 48 herbaceous understory species and estimated small-scale species richness in early and late successional boreonemoral herb-rich coniferous forests in central Estonia. VM of erosulate growth forms was significantly higher than that of hemi-rosette and rosette growth forms. Erosulate species exhibited higher mobility in young stands, but their relative and total cover was considerably higher in old stands. Local plant richness (in 1 x 1 m plots) correlated positively with the variability of VM of species in a plot--larger differences in VM resulted in a higher number of coexisting species. Our results thus suggest that species differences in VM can contribute to small-scale coexistence by providing different ways to colonise empty space.

Foraging for space and avoidance of physical obstructions by plant roots: a comparative study of grasses from contrasting habitats.

Physical obstructions that reduce space for root growth can profoundly affect plant performance. The aim of this study was to investigate the ability of roots to avoid obstructions and forage for usable space, and to reveal the mechanism involved. Eight grass species from four genera were examined. Each genus included species characteristic of habitats with high and low nutrient availability. The ability to limit root mass and to adjust morphology within substrate containing obstructions in the form of gravel was investigated. A treatment with activated carbon, which adsorbs organic compounds, was used to examine the possible involvement of root exudates in responses to obstructions. Only species characteristic of nutrient-poor habitats restricted placement of root mass in substrate containing obstructions, and this response disappeared in the presence of activated carbon. Root morphological responses to obstructions differed from those shown in response to nutrient-poor conditions or compacted soil. These results suggest that the ability to avoid obstructions is dependent on the sensitivity of roots to their own exudates accumulating in the vicinity of obstructions. This is similar to other behavioural responses in which cues or signals are used to adjust growth before stressful conditions are encountered.

Climate warming may facilitate invasion of the exotic shrub Lantana camara.

Plant species show different responses to the elevated temperatures that are resulting from global climate change, depending on their ecological and physiological characteristics. The highly invasive shrub Lantana camara occurs between the latitudes of 35 °N and 35 °S. According to current and future climate scenarios predicted by the CLIMEX model, climatically suitable areas for L. camara are projected to contract globally, despite expansions in some areas. The objective of this study was to test those predictions, using a pot experiment in which branch cuttings were grown at three different temperatures (22 °C, 26 °C and 30 °C). We hypothesized that warming would facilitate the invasiveness of L. camara. In response to rising temperatures, the total biomass of L. camara did increase. Plants allocated more biomass to stems and enlarged their leaves more at 26 °C and 30 °C, which promoted light capture and assimilation. They did not appear to be stressed by higher temperatures, in fact photosynthesis and assimilation were enhanced. Using lettuce (Lactuca sativa) as a receptor plant in a bioassay experiment, we also tested the phytotoxicity of L. camara leachate at different temperatures. All aqueous extracts from fresh leaves significantly inhibited the germination and seedling growth of lettuce, and the allelopathic effects became stronger with increasing temperature. Our results provide key evidence that elevated temperature led to significant increases in growth along with physiological and allelopathic effects, which together indicate that global warming facilitates the invasion of L. camara.

Ecological assembly rules in plant communities--approaches, patterns and prospects.

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

Limited phenotypic plasticity in range-edge populations: a comparison of co-occurring populations of two Agrimonia species with different geographical distributions.

Increased importance of genetic drift and selection for stress resistance have been predicted to lead to a reduction in the degree of phenotypic plasticity in populations at margins of a species' geographical range, relative to those in the centre. We examined the effect of population positioning within the species range on degree of active morphological plasticity to vegetation shade. Importantly, we discriminated between active, size-independent morphological adjustments in response to shade and passive changes in morphology caused by the dependence of morphological traits on plant size, as only the former are considered to be adaptive. Two closely related and ecologically similar Agrimonia species were examined in the same geographical location, where one species reaches the edge of its distribution (Agrimonia pilosa) and the other does not (A. eupatoria). Plasticity to light availability is likely to be advantageous for both species as they occupy habitats with variable light conditions. However, we hypothesised that high levels of environmental stress should lead to reduced active plasticity in marginal compared with more central populations. Agrimonia eupatoria exhibited active adjustments in leaf morphology in response to tree shade, and in elongation of stems and inflorescences in response to herbaceous shade. In contrast, A. pilosa exhibited very limited active plasticity. High active plasticity allowed A. eupatoria to retain constant shoot growth in a wide range of light conditions, while the lack of active plasticity in A. pilosa resulted in a strong dependence of shoot growth on light availability. We propose that high levels of environmental stress in marginal areas of a species' range may lead to a significant reduction in the degree of active plasticity. Our results clearly indicate that discrimination between active and passive plasticity is crucial for reaching valid conclusions about differences in adaptive plasticity between marginal and non-marginal populations.

The role of leaf lobation in elongation responses to shade in the rosette-forming forb Serratula tinctoria (Asteraceae).

BACKGROUND AND AIMS: Lobed leaves are considered selectively advantageous in conditions of high irradiance. However, most studies have involved woody species, with only a few considering the role of leaf lobation in herbaceous rosette species. In this study, it is hypothesized that, in addition to its adaptive value in high light, leaf lobation may add to the function of petioles as vertical spacers in herbaceous species in conditions of strong competition for light. METHODS: To test this hypothesis, leaf development was examined under seasonally changing natural light conditions and a field experiment was conducted in which light climate was manipulated in a wooded meadow population of Serratula tinctoria. KEY RESULTS: No changes in leaf lobation were observed in response to experimental shading or different natural light conditions. However, in tall herbaceous vegetation, plants with highly lobed leaves achieved significantly greater vertical elongation than plants with less-lobed leaves. In contrast to herbaceous shade, tree shade had no effect on leaf elongation, suggesting differential responsiveness to competition from neighbouring herbs versus overhead shade. In shading treatments, imposed shade could only be responded to by the elongation of leaves that were produced late in development. CONCLUSIONS: The results show that extensive leaf lobation can enable greater leaf elongation in response to shade from surrounding herbaceous vegetation. The different morphological responses displayed by Serratula tinctoria to different types of shade demonstrate the importance of critically assessing experimental designs when investigating phenotypic plasticity in response to shade.

Monitoring of biological diversity: a common-ground approach.

Practical approaches to monitoring biological diversity vary widely among countries, and the accumulating data are frequently not generalizable at the international scale. Although many present monitoring schemes, especially in developed countries, produce highly complex data, there is often a lack of basic data about the level and spatial distribution of biodiversity. We augmented the general framework for improving biomonitoring, proposed by Green et al. (2005), and identified its core tasks and attributes. The first priority for a more unified biodiversity monitoring is to agree on a minimum set of core tasks and attributes, which will make it possible to build a standardized biomonitoring system even in countries with few resources. Our scheme has two main organizational levels-taxa and ecosystems. The basic elements of the biomonitoring system proposed are recording of presence and absence of taxa and ecosystems in a target area, mapping of their distribution in space, and assessment of their status. All the elements have to be repeated over time. Although these tasks are fundamental, they are frequently not considered in currently functioning biomonitoring programs. The whole system has to be hierarchical and additive: if more resources are available, new activities may be added to the basic routine. Agreeing on a common standard will facilitate aggregating measures of biodiversity status and trends into regional and global indices. This information will relate directly to several Convention on Biological Diversity indicators for assessing progress toward the 2010 Biodiversity Target.

High shoot plasticity favours plant coexistence in herbaceous vegetation.

Several theoretical considerations imply that high shoot morphological plasticity could increase competition symmetry and favour plant coexistence. We tested whether mean plasticity across co-occurring species is a key trait for explaining ramet density and species richness in herbaceous vegetation. We used three data sets to test the hypotheses: (a) experimentally achieved estimates of plasticity to light availability for 35 herbaceous species; (b) richness, ramet density and canopy architecture data from 17 herbaceous communities; (c) species richness data from a 5-year permanent-plot study in a calcareous grassland. In herbaceous communities containing species with relatively higher shoot plasticity, ramet density was significantly higher. Consequently, relatively more species were growing per unit area-a greater proportion of the community species pool was represented on 1 m(2). In the permanent plot study species-richness was higher in those 40 x 40 cm quadrats where species with high shoot plasticity prevailed-there was a positive regression of richness on the mean plasticity of species. This relationship was highly significant in five consecutive years. Our results suggest that shoot plasticity to light availability is evidently one of the key traits in processes that alter the density of co-existing plants and, therefore, species diversity in herbaceous communities.