Ecological restoration outcomes are highly variable, undermining efforts to recover biodiversity and ecosystem functions. One poorly understood source of variability is ‘year effects’—interannual variation in environmental conditions during the first year of restoration that alter successional trajectories of plant communities. There have been few experimental tests disentangling planting years from other differences among restoration projects (e.g. edaphic conditions, restoration approach), particularly those resolving mechanisms for year effects such as planting‐year rainfall. Moreover, past year effect studies focused almost exclusively on species‐level consequences. Therefore, the extent to which year effects influence the traits of communities is unknown. To address these gaps and provide a mechanistic test of how precipitation contributes to year effects, we conducted an experiment where we manipulated rainfall (drought, average and high levels) during the first growing season, replicated across three establishment year treatments to disentangle the effects of precipitation from other drivers of year effects. In each establishment year, we seeded the same species mix to initiate grassland restoration. We then surveyed plant community compositions annually for 5 years to quantify trait responses of restored communities to planting year rainfall. We found that variation in planting‐year precipitation altered community assembly trajectories by influencing community‐weighted mean (CWM) trait composition, and these effects persisted for at least 5 years. Over time, CWM specific leaf area and CWM seed mass decreased and CWM plant height increased. The effect of age on CWM plant height was stronger in plots that received mean and high watering treatments compared to drought treatments. This effect was also observed for CWM seed mass, albeit weaker. We also found some evidence for planting year effects unrelated to planting‐year rainfall for the three CWM traits, illustrating how interannually varying environmental conditions besides rainfall can generate persistent year effect on plant communities through their traits.
Despite theoretical advances, the ecological factors and functional traits that enable species varying in seed size and fecundity to coexist remain unclear. Given inherent fecundity advantages, why don't small‐seeded species dominate communities? In perennial grasslands, we evaluated whether small‐seeded species are less tolerant of competition from the community dominant bunchgrass than large‐seeded species, but also less vulnerable to seed predation by mice. We also explored whether trade‐offs involving competitive tolerance include two other functional traits, height and leaf mass per area (LMA). We added seeds of 17 forb species to plots where bunchgrass competition and rodent seed predation were manipulated across sites varying in bunchgrass productivity and thus competitive intensity. Seeds were added at densities mimicking interspecific variation in fecundity among target species. Standardizing for differences in fecundity (i.e. seed input, which enabled us to evaluate inherent interspecific differences in susceptibility to biotic interactions), bunchgrass competition more greatly reduced recruitment and establishment of small‐ versus large‐seeded species, whereas rodent seed predation more greatly reduced the recruitment of large‐ versus small‐seeded species. Plant height and LMA were unrelated to the competition effect size. Small‐seeded species abundance decreased across sites increasing in bunchgrass productivity, whereas this was not the case for large‐seeded species. For adult plants but not seedlings, community‐weighted functional trait means (CWM) for seed size, height and LMA increased in plots with versus without bunchgrass competition and the CWM for seed size and height also increased at sites with greater bunchgrass productivity (for adults only). In contrast, rodent seed predation had no significant effects on CWM seed size. At the end of the experiment, adult abundance positively correlated with plant fecundity in plots lacking bunchgrass, indicating the inherent advantages accrued to high fecundity small‐seeded species. However, with bunchgrass competition, abundances were equalized across species due to reduced competitive tolerance of high fecundity small‐seeded species.
- NSF-PAR ID:
- 10379064
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Journal of Ecology
- Volume:
- 109
- Issue:
- 2
- ISSN:
- 0022-0477
- Page Range / eLocation ID:
- p. 626-638
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
More Like this
-
more » « less
Abstract Synthesis and applications . Our results provide evidence for planting year rainfall interacting with community assembly to alter the functional trait composition of restored grasslands. This suggests that interannual variation in rainfall during establishment is an important source of divergent biodiversity and functional outcomes in restored grasslands. -
more » « less
Abstract Grasslands are subject to climate change, such as severe drought, and an important aspect of their functioning is temporal stability in response to extreme climate events. Previous research has explored the impacts of extreme drought and post‐drought periods on grassland stability, yet the mechanistic pathways behind these changes have rarely been studied.
Here, we implemented an experiment with 4 years of drought and 3 years of recovery to assess the effects of drought and post‐drought on the temporal stability of above‐ground net primary productivity (ANPP) and its underlying mechanisms. To do so, we measured community‐weighted mean (CWM) of six plant growth and nine seed traits, functional diversity, population stability and species asynchrony across two cold, semiarid grasslands in northern China. We also performed piecewise structural equation models (SEMs) to assess the relationships between ANPP stability and its underlying mechanisms and how drought and post‐drought periods alter the relative contribution of these mechanisms to ANPP stability.
We found that temporal stability of ANPP was not reduced during drought due to grasses maintaining productivity, which compensated for increased variation of forb productivity. Moreover, ANPP recovered rapidly after drought, and both grasses and forbs contributed to community stability during the post‐drought period. Overall, ANPP stability decreased during the combined drought and post‐drought periods because of rapid changes in ANPP from drought to post‐drought. SEMs revealed that the temporal stability of ANPP during drought and post‐drought periods was modulated by functional diversity and community‐weighted mean traits directly and indirectly by altering species asynchrony and population stability. Specifically, the temporal stability of ANPP was positively correlated with functional divergence of plant communities. CWMs of seed traits (e.g. seed width and thickness), rather than plant growth traits (e.g. specific leaf area and leaf nutrient content), stabilized grassland ANPP. Productivity of plant communities with large and thick seeds was less sensitive to precipitation changes over time.
These results emphasize the importance of considering both the functional trait distribution among species and seed traits of dominant species since their combined effects can stabilize ecosystem functions under global climate change scenarios.
Read the free
Plain Language Summary for this article on the Journal blog. -
more » « less
Abstract Traits differentially adapt plant species to particular conditions generating compositional shifts along environmental gradients. As a result, community‐scale trait values show concomitant shifts, termed trait‒environment relationships. Trait‒environment relationships are often assessed by evaluating community‐weighted mean (CWM) traits observed along environmental gradients. Regression‐based approaches (CWMr) assume that local communities exhibit traits centred at a single optimum value and that traits do not covary meaningfully. Evidence suggests that the shape of trait‒abundance relationships can vary widely along environmental gradients—reflecting complex interactions—and traits are usually interrelated. We used a model that accounts for these factors to explore trait‒environment relationships in herbaceous forest plant communities in Wisconsin (USA).
We built a generalized linear mixed model (GLMM) to analyse how abundances of 185 species distributed among 189 forested sites vary in response to four functional traits (vegetative height—VH, leaf size—LS, leaf mass per area—LMA and leaf carbon content), six environmental variables describing overstorey, soil and climate conditions, and their interactions. The GLMM allowed us to assess the nature and relative strength of the resulting 24 trait‒environment relationships. We also compared results between GLMM and CWMr to explore how conclusions differ between approaches.
The GLMM identified five significant trait‒environment relationships that together explain ~40% of variation in species abundances across sites. Temperature appeared as a key environmental driver, with warmer and more seasonal sites favouring taller plants. Soil texture and temperature seasonality affected LS and LMA; seasonality effects on LS and LMA were nonlinear, declining at more seasonal sites. Although often assumed for CWMr, only some traits under certain conditions had centred optimum trait‒abundance relationships. CWMr more liberally identified (13) trait‒environment relationships as significant but failed to detect the temperature seasonality‒LMA relationship identified by the GLMM.
Synthesis . Although GLMM represents a more methodologically complex approach than CWMr, it identified a reduced set of trait‒environment relationships still capable of accounting for the responses of forest understorey herbs to environmental gradients. It also identified separate effects of mean and seasonal temperature on LMA that appear important in these forests, generating useful insights and supporting broader application of GLMM approach to understand trait‒environment relationships. -
more » « less
Abstract Body size influences an individual's physiology and the nature of its intra‐ and interspecific interactions. Changes in this key functional trait can therefore have important implications for populations as well. For example, among invertebrates, there is typically a positive correlation between female body size and reproductive output. Increasing body size can consequently trigger changes in population density, population structure (e.g. adult to juvenile ratio) and the strength of intraspecific competition.
Body size changes have been documented in several species in the Arctic, a region that is warming rapidly. In particular, wolf spiders, one of the most abundant arctic invertebrate predators, are becoming larger and therefore more fecund. Whether these changes are affecting their populations and role within food webs is currently unclear.
We investigated the population structure and feeding ecology of the dominant wolf spider species
Pardosa lapponica at two tundra sites where adult spiders naturally differ in mean body size. Additionally, we performed a mesocosm experiment to investigate how variation in wolf spider density, which is likely to change as a function of body size, influences feeding ecology and its sensitivity to warming.We found that juvenile abundance is negatively associated with female size and that wolf spiders occupied higher trophic positions where adult females were larger. Because female body size is positively related to fecundity in
P. lapponica , the unexpected finding of fewer juveniles with larger females suggests an increase in density‐dependent cannibalism as a result of increased intraspecific competition for resources. Higher rates of density‐dependent cannibalism are further supported by the results from our mesocosm experiment, in which individuals occupied higher trophic positions in plots with higher wolf spider densities. We observed no changes in wolf spider feeding ecology in association with short‐term experimental warming.Our results suggest that body size variation in wolf spiders is associated with variation in intraspecific competition, feeding ecology and population structure. Given the widespread distribution of wolf spiders in arctic ecosystems, body size shifts in these predators as a result of climate change could have implications for lower trophic levels and for ecosystem functioning.
-
more » « less
Abstract Under fire suppression, many tropical savannas transform into forests. Forest expansion entails changes in environmental variables and plant community structure. We hypothesized that forest expansion into savanna results in a shift in community‐weighted mean functional traits from stress tolerance to competitiveness, with generalist species having trait values intermediate between those of specialists of savanna and forest habitats.
We studied 30 plots distributed over three savanna–forest boundaries undergoing forest expansion in the Brazilian Cerrado, capturing a gradient from open savanna to recently formed forest. We measured functional traits of 116 woody species of savanna specialist, generalist and forest specialist functional groups and quantified changes in species composition and mean traits across the basal area gradient.
We identified two main axes of species traits. The first separated forest and generalist species from savanna specialists, with the latter possessing traits associated with resistance to disturbance and stress— such as thick leaves, thick bark, slower height growth and lower shade tolerance. Our second trait axis separated shrubs and understorey trees from pioneer species. Generalist species’ traits did not differ substantially from forest species, nor did they tend to have a typical pioneer strategy.
Community‐weighted trait means changed linearly with forest development. There was a steady increase in traits associated with competitive dominance rather than stress tolerance and fire resistance, indicating a wholesale shift in the selective environment. Several of these patterns—for example, increasing height and decreasing light requirements—are common in old‐field succession. In contrast to old‐field succession, we found that SLA increased, leaf thickness decreased and wood density stayed constant.
The assembly of forests appears to be shaped by environmental filters that contribute to a functional trajectory distinct from most other studied ecosystems. Our results highlight the importance of the functional composition of the early community and of the early colonizers of the open environment. Differences between savanna and forest specialists reflect the selective effects of the contrasting environments, while the traits of generalists—and their interaction with environmental filters—drive the dynamics of forest expansion.
A free
Plain Language Summary can be found within the Supporting Information of this article.
