Abstract. Over the last century, many grasslands worldwide have transitioned from a graminoid to a tree/shrub-dominated state in a short period of time, a phenomenon referred to as woody encroachment. Positive feedbacks and bi-stability are thought to be important drivers of woody encroachment, but there is little empirical evidence to suggest that positive feedbacks accelerate the woody encroachment of mesic grasslands. In mesic tallgrass prairie, shrub establishment does not directly facilitate seedling establishment. Yet, shrub establishment may facilitate the clonal spread of existing shrubs into nearby patches, because clonal reproduction might circumvent barriers that typically limit woody seedlings. Our results show that when Cornus drummondii (the predominate encroacher of mesic tallgrass prairie) extends rhizomatous stems into open grasslands, these stems use the same deep soil water sources as mature stems-thereby avoiding competition with grasses and gaining access to a reliable water source. In addition, herbaceous fuel concentrations are lower at the shrub/grass interface than in open grasslands, reducing the potential impacts of subsequent grassland fires. We propose that the release from resource and fire limitation results in a positive feedback loop as clonal stems are able to extend into surrounding patches, circumvent demographic barriers, mature, and spread by developing their own clonal stems. Long-term data on site (26 years) corroborates this interpretation: the size of deep-rooted clonal shrub species has increased 16-fold and their cover has increased from 0 to 27%, whereas the cover of shallowrooted species (both clonal and non-clonal) has only increased marginally. Together, these results suggest that (1) positive feedbacks can facilitate mesic woody encroachment and (2) bi-stability exists in mesic tallgrass prairie.
Plant traits can provide unique insights into plant performance at the community scale. Functional composition, defined by both functional diversity and community‐weighted trait means (CWMs), can affect the stability of above‐ground net primary production (ANPP) in response to climate extremes. Further complexity arises, however, when functional composition itself responds to environmental change. The duration of climate extremes, such as drought, is expected to increase with rising global temperatures; thus, understanding the impacts of long‐term drought on functional composition and the corresponding effect that has on ecosystem function could improve predictions of ecosystem sensitivity to climate change. We experimentally reduced growing season precipitation by 66% across six temperate grasslands for 4 years and measured changes in three indices of functional diversity (functional dispersion, richness and evenness), community‐weighted trait means and phylogenetic diversity (PD). Specific leaf area (SLA), leaf nitrogen content (LNC) and (at most sites) leaf turgor loss point (πTLP) were measured for species cumulatively representing ~90% plant cover at each site. Long‐term drought led to increased community functional dispersion in three sites, with negligible effects on the remaining sites. Species re‐ordering following the mortality/senescence of dominant species was the main driver of increased functional dispersion. The response of functional diversity was not consistently matched by changes in phylogenetic diversity. Community‐level drought strategies (assessed as CWMs) largely shifted from drought tolerance to drought avoidance and/or escape strategies, as evidenced by higher community‐weighted πTLP, SLA and LNC. Lastly, ecosystem drought sensitivity (i.e. relative reduction in ANPP in drought plots) was positively correlated with community‐weighted SLA and negatively correlated with functional diversity. Synthesis. Increased functional diversity following long‐term drought may stabilize ecosystem functioning in response to future drought. However, shifts in community‐scale drought strategies may increase ecosystem drought sensitivity, depending on the nature and timing of drought. Thus, our results highlight the importance of considering both functional diversity and abundance‐weighted traits means of plant communities as their collective effect may either stabilize or enhance ecosystem sensitivity to drought.
Plant traits can be used to predict ecosystem responses to environmental change using a response–effect trait framework. To do this, appropriate traits must be identified that explain a species' influence on ecosystem function (“effect traits”) and the response of those species to environmental change (“response traits”). Response traits are often identified and measured along gradients in plant resources, such as water availability; however, precipitation explains very little variation in most plant traits globally. Given the strong relationship between plant traits and ecosystem functions, such as net primary productivity (NPP), and between NPP and precipitation, the lack of correlation between precipitation and plant traits is surprising. We address this issue through a systematic review of >500 published studies that describe plant trait responses to altered water availability. The overarching goal of this review was to identify potential causes for the weak relationship between commonly measured plant traits and water availability so that we may identify more appropriate “response traits.” We attribute weak trait–precipitation relationships to an improper selection of traits (e.g., nonhydraulic traits) and a lack of trait‐based approaches that adjust for trait variation within communities (only 4% of studies measure community‐weighted traits). We then highlight the mechanistic value of hydraulic traits as more appropriate “response traits” with regard to precipitation, which should be included in future community‐scale trait surveys. Trait‐based ecology has the potential to improve predictions of ecosystem responses to predicted changes in precipitation; however, this predictive power depends heavily on the identification of reliable response and effect traits. To this end, trait surveys could be improved by a selection of traits that reflect physiological functions directly related to water availability with traits weighted by species relative abundance. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13135/suppinfo is available for this article.
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