Summary 1. Chronosequences and associated space-for-time substitutions are an important and often necessary tool for studying temporal dynamics of plant communities and soil development across multiple time-scales. However, they are often used inappropriately, leading to false conclusions about ecological patterns and processes, which has prompted recent strong criticism of the approach. Here, we evaluate when chronosequences may or may not be appropriate for studying community and ecosystem development. 2. Chronosequences are appropriate to study plant succession at decadal to millennial time-scales when there is evidence that sites of different ages are following the same trajectory. They can also be reliably used to study aspects of soil development that occur between temporally linked sites over time-scales of centuries to millennia, sometimes independently of their application to shorter-term plant and soil biological communities. 3. Some characteristics of changing plant and soil biological communities (e.g. species richness, plant cover, vegetation structure, soil organic matter accumulation) are more likely to be related in a predictable and temporally linear manner than are other characteristics (e.g. species composition and abundance) and are therefore more reliably studied using a chronosequence approach. 4. Chronosequences are most appropriate for studying communities that are following convergent successional trajectories and have low biodiversity, rapid species turnover and low frequency and severity of disturbance. Chronosequences are least suitable for studying successional trajectories that are divergent, species-rich, highly disturbed or arrested in time because then there are often major difficulties in determining temporal linkages between stages. 5. Synthesis. We conclude that, when successional trajectories exceed the life span of investigators and the experimental and observational studies that they perform, temporal change can be successfully explored through the judicious use of chronosequences.

/pdf/the-use-of-chronosequences-in-studies-of-ecological-1c5s5077we.pdf

The use of chronosequences in studies of ecological succession and soil development

Tropical forest vegetation is shaped by climate and by soil, but understanding how the distributions of individual tree species respond to specific resources has been hindered by high diversity and consequent rarity. To study species over an entire community, we surveyed trees and measured soil chemistry across climatic and geological gradients in central Panama and then used a unique hierarchical model of species occurrence as a function of rainfall and soil chemistry to circumvent analytical difficulties posed by rare species. The results are a quantitative assessment of the responses of 550 tree species to eight environmental factors, providing a measure of the importance of each factor across the entire tree community. Dry-season intensity and soil phosphorus were the strongest predictors, each affecting the distribution of more than half of the species. Although we anticipated clear-cut responses to dry-season intensity, the finding that many species have pronounced associations with either high or low phosphorus reveals a previously unquantified role for this nutrient in limiting tropical tree distributions. The results provide the data necessary for understanding distributional limits of tree species and predicting future changes in forest composition.

/pdf/species-distributions-in-response-to-individual-soil-1ptzjm8iy1.pdf

Species distributions in response to individual soil nutrients and seasonal drought across a community of tropical trees

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.

A guide for using functional diversity indices to reveal changes in assembly processes along ecological gradients

Lessons learned from the study of ecological succession have much to offer contemporary environmental problem solving but these lessons are being underutilized. As anthropogenic disturbances increase, succession is more relevant than ever. In this review, we suggest that succession is particularly suitable to address concerns about biodiversity loss, climate change, invasive species, and ecological restoration. By incorporating modern experimental techniques and linking results across environmental gradients with meta-analyses, studies of succession can substantially improve our understanding of other ecological phenomena. Succession can help predict changes in biodiversity and ecosystem services impacted by invasive species and climate change and guide manipulative responses to these disruptions by informing restoration efforts. Succession is still a critical, integrative concept that is central to ecology.

https://www.prf.jcu.cz/files/news/PrachWalker2011.pdf

Four opportunities for studies of ecological succession

Summary 1. Theory predicts that the processes generating biodiversity after disturbance will change during succession. Comparisons of phylogenetic and functional (alpha and beta) diversity with taxonomic diversity can provide insights into the extent to which community assembly is driven by deterministic or stochastic processes, but comparative approaches have yet to be applied to successional systems. 2. We characterized taxonomic, phylogenetic and functional plant (alpha and beta) diversity within and between four successional stages in a > 270-year-long arable-to-grassland chronosequence. Null models were used to test whether functional and phylogenetic turnover differed from random expectations, given the levels of species diversity. 3. The three facets of diversity showed different patterns of change during succession. Between early and early-mid succession, species richness increased but there was no increase in functional or phylogenetic diversity. Higher than predicted levels of functional similarity between species within the early and early-mid successional stages, indicate that abiotic filters have selected for sets of functionally similar species within sites. Between late-mid and late succession, there was no further increase in species richness, but a significant increase in functional alpha diversity, suggesting that functionally redundant species were replaced by functionally more dissimilar species. Functional turnover between stages was higher than predicted, and higher than within-stage turnover, indicating that different assembly processes act at different successional stages. 4. Synthesis. Analysis of spatial and temporal turnover in different facets of diversity suggests that deterministic processes generate biodiversity during post-disturbance ecosystem development and that the relative importance of assembly processes has changed over time. Trait-mediated abiotic filtering appears to play an important role in community assembly during the early and early-mid stages of arable-to-grassland succession, whereas the relative importance of competitive exclusion appears to have increased towards the later successional stages. Phylogenetic diversity provided a poor reflection of functional diversity and did not contribute to inferences about underlying assembly processes. Functionally deterministic assembly suggests that it may be possible to predict future post-disturbance changes in biodiversity, and associated ecosystem attributes, on the basis of species’ functional traits but not phylogeny.

/pdf/contrasting-changes-in-taxonomic-phylogenetic-and-functional-t483wkzotl.pdf

Contrasting changes in taxonomic, phylogenetic and functional diversity during a long‐term succession: insights into assembly processes

Over time scales of thousands to millions of years, and in the absence of rejuvenating disturbances that initiate primary or early secondary succession, ecosystem properties such as net primary productivity, decomposition, and rates of nutrient cycling undergo substantial declines termed ecosystem retrogression. Retrogression results from the depletion or reduction in the availability of nutrients, and can only be reversed through rejuvenating disturbance that resets the system; this differs from age-related declines in forest productivity that are driven by shorter-term depression of nutrient availability and plant ecophysiological process rates that occur during succession. Here we review and synthesize the findings from studies of long-term chronosequences that include retrogressive stages for systems spanning the boreal, temperate, and subtropical zones. Ecosystem retrogression has been described by ecologists, biogeochemists, geologists, and pedologists, each of which has developed somewhat independent conceptual frameworks; our review seeks to unify this literature in order to better understand the causes and consequences of retrogression. Studies of retrogression have improved our knowledge of how long-term pedogenic changes drive shorter-term biological processes, as well as the consequences of these changes for ecosystem development. Our synthesis also reveals that similar patterns of retrogression (involving reduced soil fertility, predictable shifts in organismic traits, and ecological processes) occur in systems with vastly different climatic regimes, geologic substrates, and vegetation types, even though the timescales and mechanisms driving retrogression may vary greatly among sites. Studies on retrogression also provide evidence that in many regions, high biomass or "climax" forests are often transient, and do not persist indefinitely in the absence of rejuvenating disturbance. Finally, our review highlights that studies on retrogressive chronosequences in contrasting regions provide unparalleled opportunities for developing general principles about the long-term feedbacks between biological communities and pedogenic processes, and how these control ecosystem development.

W. Troy Baisden

Papers

Understanding ecosystem retrogression