Plant clonality in a soil-impoverished open ecosystem: insights from southwest Australian shrublands.

BACKGROUND AND AIMS: Clonality is a key life-history strategy promoting on-spot persistence, space occupancy, resprouting after disturbance, and resource storage, sharing and foraging. These functions provided by clonality can be advantageous under different environmental conditions, including resource-paucity and fire-proneness, which define most mediterranean-type open ecosystems, such as southwest Australian shrublands. Studying clonality-environment links in underexplored mediterranean shrublands could therefore deepen our understanding of the role played by this essential strategy in open ecosystems globally. METHODS: We created a new dataset including 463 species, six traits related to clonal growth organs (CGOs; lignotubers, herbaceous and woody rhizomes, stolons, tubers, stem fragments), and edaphic predictors of soil water availability, nitrogen (N) and phosphorus (P) from 138 plots. Within two shrubland communities, we explored multivariate clonal patterns and how the diversity of CGOs, and abundance-weighted and unweighted proportions .of clonality in plots changed along with the edaphic gradients. KEY RESULTS: We found clonality in 65 % of species; the most frequent were those with lignotubers (28 %) and herbaceous rhizomes (26 %). In multivariate space, plots clustered into two groups, one distinguished by sandy plots and plants with CGOs, the other by clayey plots and non-clonal species. CGO diversity did not vary along the edaphic gradients (only marginally with water availability). The abundance-weighted proportion of clonal species increased with N and decreased with P and water availability, yet these results were CGO-specific. We revealed almost no relationships for unweighted clonality. CONCLUSIONS: Clonality is more widespread in shrublands than previously thought, and distinct plant communities are distinguished by specific suites (or lack) of CGOs. We show that weighting belowground traits by aboveground abundance affects the results, with implications for trait-based ecologists using abundance-weighting. We suggest unweighted approaches for belowground organs in open ecosystems until belowground abundance is quantifiable.

Incorporating clonality into the plant ecology research agenda.

A longstanding research divide exists in plant ecology: either focusing on plant clonality, with no ambition to address nonclonal plants, or focusing on all plants, ignoring that many ecological processes can be affected by the fact that some plants are clonal while others are not. This gap cascades into a lack of distinction and knowledge about the similarities and differences between clonal and nonclonal plants. Here we aim to bridge this gap by identifying areas that would benefit from the incorporation of clonal growth into one integrated research platform: namely, response to productivity and disturbance, biotic interactions, and population dynamics. We are convinced that this will provide a roadmap to gain valuable insights into the ecoevolutionary dynamics relevant to all plants.

Strong impact of management regimes on rhizome biomass across Central European temperate grasslands.

Grassland ecosystems account for approximately 40% of terrestrial biomes globally. These communities are characterized by a large allocation to belowground biomass, often exceeding its aboveground counterpart. However, this biomass investment cannot be entirely attributed to the acquisitive function of roots. Grassland plants also allocate to non-acquisitive, stem-derived, belowground organs, such as rhizomes. These organs are responsible for the key plant functions of space occupancy, resprouting after damage, and seasonal rest. However, biomass investment to rhizomes has rarely been studied. Here we gathered community-level aboveground and rhizome biomass data for 52 temperate grasslands in Czech Republic (Central Europe), differing in management intensity. We found that rhizome biomass scaled linearly with aboveground biomass, and more intensive management disproportionally (negatively) affected rhizome biomass. This finding may have important implications for the persistence of temperate grassland plants and their provision of ecosystem services (e.g., soil carbon sequestration, soil stabilization) in relation to changing environments.

The Neglected Belowground Dimension of Plant Dominance.

Dominants are key species that shape ecosystem functioning. Plant dominance is typically assessed on aboveground features. However, belowground, individual species may not scale proportionally in relation to their aboveground dimension. This is especially important in ecosystems where most biomass is allocated belowground, including grassy and shrubby biomes.

Linking Plant Functional Ecology to Island Biogeography.

The study of insular systems has a long history in ecology and biogeography. Island plants often differ remarkably from their noninsular counterparts, constituting excellent models for exploring eco-evolutionary processes. Trait-based approaches can help to answer important questions in island biogeography, yet plant trait patterns on islands remain understudied. We discuss three key hypotheses linking functional ecology to island biogeography: (i) plants in insular systems are characterized by distinct functional trait syndromes (compared with noninsular environments); (ii) these syndromes differ between true islands and terrestrial habitat islands; and (iii) island characteristics influence trait syndromes in a predictable manner. We are convinced that implementing trait-based comparative approaches would considerably further our understanding of plant ecology and evolution in insular systems.

Community weighted mean trait data of Italian forest understories.

Plant functional trait data aggregated at the community level (i.e., community weighted mean, CWM) are fundamental to study plant-environment relationships. Here, we provide a large database of CWM values of twelve traits reflecting several plant functions, including leaf, seed, whole-plant, clonal and bud bank traits. The CWMs were calculated in 201 forest stands (a statistically representative sample of all the Italian forests) across three biogeographic regions: Alpine, Continental, and Mediterranean.

Towards an eco-evolutionary understanding of endemism hotspots and refugia.

Background: Refugia are island-like habitats that are linked to long-term environmental stability and, as a result, high endemism. Conservation of refugia and endemism hotspots should be based on a deep ecological and evolutionary understanding of their functioning, which remains limited. Although functional traits can provide such insights, a corresponding, coherent framework is lacking. Proposed Framework: Plant communities in refugia and endemism hotspots should, due to long-term environmental stability, display unique functional characteristics linked to distinct phylogenetic patterns. Therefore, such communities should be characterized by a functional signature that exhibits: (1) distinct values and combinations of traits, (2) higher functional diversity and (3) a prevalence of similar traits belonging to more distantly related lineages inside, compared to outside, of endemism hotspots and refugia. While the limited functional trait data available from refugia and endemism hotspots do not allow these predictions to be tested rigorously, three potential applications of the functional signature in biogeography and conservation planning are highlighted. Firstly, it allows the functional characteristics of endemism hotspots and refugia to be identified. Secondly, the strength of the functional signature can be compared among these entities, and with the surrounding landscape, to provide an estimate of the capacity of endemism hotspots and refugia to buffer environmental changes. Finally, the pattern of the functional signature can reveal ecological and evolutionary processes driving community assembly and functioning, which can assist in predicting the effect of environmental changes (e.g. climate, land-use) on communities in endemism hotspots and refugia. Conclusion: The proposed functional signature concept allows the systematic integration of plant functional traits and phylogeny into the study of endemism hotspots and refugia, but more data on functional traits in these entities are urgently needed. Overcoming this limitation would facilitate rigorous testing of the proposed predictions for the functional signature, advancing the eco-evolutionary understanding of endemism hotspots and refugia.

Quantifying the effects of ecological constraints on trait expression using novel trait-gradient analysis parameters.

Complex processes related to biotic and abiotic forces can impose limitations to assembly and composition of plant communities. Quantifying the effects of these constraints on plant functional traits across environmental gradients, and among communities, remains challenging. We define ecological constraint (Ci ) as the combined, limiting effect of biotic interactions and environmental filtering on trait expression (i.e., the mean value and range of functional traits). Here, we propose a set of novel parameters to quantify this constraint by extending the trait-gradient analysis (TGA) methodology. The key parameter is ecological constraint, which is dimensionless and can be measured at various scales, for example, on population and community levels. It facilitates comparing the effects of ecological constraints on trait expressions across environmental gradients, as well as within and among communities. We illustrate the implementation of the proposed parameters using the bark thickness of 14 woody species along an aridity gradient on granite outcrops in southwestern Australia. We found a positive correlation between increasing environmental stress and strength of ecological constraint on bark thickness expression. Also, plants from more stressful habitats (shrublands on shallow soils and in sun-exposed locations) displayed higher ecological constraint for bark thickness than plants in more benign habitats (woodlands on deep soils and in sheltered locations). The relative ease of calculation and dimensionless nature of Ci allow it to be readily implemented at various scales and make it widely applicable. It therefore has the potential to advance the mechanistic understanding of the ecological processes shaping trait expression. Some future applications of the new parameters could be investigating the patterns of ecological constraints (1) among communities from different regions, (2) on different traits across similar environmental gradients, and (3) for the same trait across different gradient types.

On Plant Modularity Traits: Functions and Challenges.

On-spot persistence, space occupancy, and recovery after damage are key plant functions largely understudied. Traits relevant to these functions are difficult to assess because of their relationships to plant modularity. We suggest that developing collection protocols for these traits is feasible and could facilitate their inclusion in global syntheses.