The study of positive species interactions is a rapidly evolving field in ecology. Despite decades of research, controversy has emerged as to whether positive and negative interactions predictably shift with increasing environmental stress as hypothesised by the stress-gradient hypothesis (SGH). Here, we provide a synthesis of 727 tests of the SGH in plant communities across the globe to examine its generality across a variety of ecological factors. Our results show that plant interactions change with stress through an outright shift to facilitation (survival) or a reduction in competition (growth and reproduction). In a limited number of cases, plant interactions do not respond to stress, but they never shift towards competition with stress. These findings are consistent across stress types, plant growth forms, life histories, origins (invasive vs. native), climates, ecosystems and methodologies, though the magnitude of the shifts towards facilitation with stress is dependent on these factors. We suggest that future studies should employ standardised definitions and protocols to test the SGH, take a multi-factorial approach that considers variables such as plant traits in addition to stress, and apply the SGH to better understand how species and communities will respond to environmental change.

https://www.onlinelibrary.wiley.com/doi/pdf/10.1111/ele.12080

Global shifts towards positive species interactions with increasing environmental stress

Salt marsh morphology is known to be strongly correlated to vegetation patterns through a complex interplay of biological and physical processes. This paper presents the results of field surveys at several study salt marshes within the Venice Lagoon (Italy), which indicate that salt-marsh macrophyte species may indeed be associated with narrow ranges of soil topographic elevation. Statistical analyses show that several properties of the frequency distributions of halophytes presence are sensitive not only to variations in soil elevation, but also to the specific marsh considered. Through direct in situ sampling and by use of a finite-element hydrodynamic model the role of plant submersion duration and frequency in determining the observed variability of vegetation species is then studied. Measurements of soil salinity have also been performed at selected salt marshes to address its influence on vegetation occurrence. With implications for tidal marshes in general, the distribution of halophytes in the salt marshes considered is found not to be responding to simple rules dictated by the tidal cycle or to salinity, and that such factors, when singularly considered, cannot explain the observed spatial distribution of halophytes. On the basis of observations and modelling results it is thus concluded that a combination of multiple factors, likely dominated by saturated/unsaturated flow in the soil, may be responsible for the observed macrophyte distribution.

/pdf/tidal-regime-salinity-and-salt-marsh-plant-zonation-1vj3ab771e.pdf

Tidal regime, salinity and salt marsh plant zonation

Mapping salt-marsh vegetation by multispectral and hyperspectral remote sensing

Underwater photosynthesis in flooded terrestrial plants: a matter of leaf plasticity.

Marshes display impressive biogeomorphic features, such as zonation, a mosaic of extensive vegetation patches of rather uniform composition, exhibiting sharp transitions in the presence of extremely small topographic gradients. Although generally associated with the accretion processes necessary for marshes to keep up with relative sea level rise, competing environmental constraints, and ecologic controls, zonation is still poorly understood in terms of the underlying biogeomorphic mechanisms. Here we find, through observations and modeling interpretation, that zonation is the result of coupled geomorphological–biological dynamics and that it stems from the ability of vegetation to actively engineer the landscape by tuning soil elevation within preferential ranges of optimal adaptation. We find multiple peaks in the frequency distribution of observed topographic elevation and identify them as the signature of biologic controls on geomorphodynamics through competing stable states modulated by the interplay of inorganic and organic deposition. Interestingly, the stable biogeomorphic equilibria correspond to suboptimal rates of biomass production, a result coherent with recent observations. The emerging biogeomorphic structures may display varying degrees of robustness to changes in the rate of sea level rise and sediment availability, with implications for the overall resilience of marsh ecosystems to climatic changes.

https://www.pnas.org/content/pnas/110/9/3259.full.pdf

Vegetation engineers marsh morphology through multiple competing stable states

The relation between vegetation zonation, elevation and inundation frequency in a Wadden Sea salt marsh

1 Zonation is often seen in environments with a strong physico-chemical gradient, such as salt marshes. It has been hypothesized that plant species are limited in their distribution by abiotic factors towards the more extreme end of the gradient, and by competition towards the more favourable end. Invasion of the native clonal grass genus Elymus in many Wadden Sea marshes may be due to increasing atmospheric nitrogen input into a nitrogen-limited environment. However, at Thmlauer Bay, Germany, Elymus athericus does not occur in lower salt marsh communities that are dominated by a dwarf shrub (Atriplex portulacoides). We therefore hypothesized that at this site the downslope (= more extreme) distributional boundary of E. athericus is a result of competition with A. portulacoides rather than of physiological limits.

2 A factorial experiment was set up to investigate the effects of removal of each competitor and fertilization. The reciprocal effects of the species on each other were measured in terms of vegetation cover and above-ground biomass. The impact of the tidal regime on plant zonation was investigated by calculating inundation frequencies at the boundary between the two plant populations from water level recordings.

3 Elymus athericus extended its distribution into the lower salt marsh when A. portulacoides was removed. The latter increased in cover but not in biomass after the removal of E. athericus. Neither species showed a response to nitrogen fertilization. The boundary between the two species in the control plots varied considerably in elevation and inundation frequency.

4 The lower distributional boundary of E. athericus can be interpreted as the result of competition with A. portulacoides. Improvement of nitrogen availability in concentrations of the same order of magnitude as annual atmospheric input had no detectable effect on plant zonation and growth. Elevation and inundation frequency were not strictly correlated with plant zonation.

Competitive exclusion of Elymus athericus from a high-stress habitat in a European salt marsh

The future perspective of salt marshes strongly depends on a positive balance between sedimentation and an increased sea-level rise. Sedimentation itself is influenced indirectly by grazing as well as the position of the location in relation to the nearest sediment source and by amount and quality of the available sediment.

Reimert Neuhaus

Papers

The relation between vegetation zonation, elevation and inundation frequency in a Wadden Sea salt marsh

Competitive exclusion of Elymus athericus from a high-stress habitat in a European salt marsh

Sedimentation in salt marshes affected by grazing regime, topographical patterns and regional differences