Thermal stress affects bioturbators' burrowing behavior: A mesocosm experiment on common cockles (Cerastoderma edule).

The intensity of marine heatwaves is increasing due to climate change. Heatwaves may affect macroinvertebrates' bioturbating behavior in intertidal areas, thereby altering the deposition-erosion balance at tidal flats. Moreover, small-scale topographic features on tidal flats can create tidal pools during the low tide, thus changing the heat capacity of tidal flats. These pools could then potentially operate as refuge environments during marine heatwaves. We studied behavior responses to heat waves using the well-known bioturbating cockle Cerastoderma edule as a model species. Different temperature regimes (i.e., fluctuating between 20 and 40 °C) and micro-topographies (i.e., presence vs. absence of tidal water pools) were mimicked in a mesocosm experiment with regular tidal regimes. Our results demonstrate that behavioral responses to heat stress strongly depend on the site-specific morphological features. Cockles covered by shallow water pools moved up when exposed to thermal stress, while burrowing deeper into the sediment in the absence of water pools. But in both cases, their migratory behavior increased under heat stress compared to regular ambient treatments. Moreover, long-term cumulative heat stress increased cockles' respiration rates and decreased their health conditions, causing mass mortality after four weeks of gradually increasing heat exposure. Overall, the present findings provide the first insights into how bioturbating behavior on tidal flats may change in response to global warming.

Modelling spatial and temporal patterns in bioturbator effects on sediment resuspension: A biophysical metabolic approach.

Tidal flats are biogeomorphic landscapes, shaped by physical forces and interaction with benthic biota. We used a metabolic approach to assess the overarching effect of bioturbators on tidal landscapes. The benthic bivalve common cockle (Cerastoderma edule) was used as model organism. The effect of C. edule on sediment resuspension was approximated as a function of the overall population metabolic rate per unit of area. We combined i) laboratory observations on how C. edule affect sediment resuspension along gradients of bioturbation activity, sediment cohesiveness and hydrodynamic force with ii) spatial data on the natural distribution of intertidal C. edule populations. This allowed us to build an integrated model of the C. edule effect on sediment resuspension along the tidal gradient. Owing to the temperature dependence of metabolic rate, the model also accounted for seasonal variation in bioturbators activity. Laboratory experiments indicated that sediment resuspension is positively related to the metabolic rate of the C. edule population especially in cohesive sediments. Based on this observation, we predicted a clear spatial and seasonal pattern in the relative importance of C. edule contribution to sediment resuspension along a tidal transect. At lower elevations, our model indicates that hydrodynamics overrules biotic effects; at higher elevations, inter-tidal hydrodynamics should be too low to suspend bioturbated sediments. The influence of C. edule on sediment resuspension is expected to be maximal at the intermediate elevation of a mudflat, owing to the combination of moderate hydrodynamic stress and high bioturbator activity. Also, bio-mediated sediment resuspension is predicted to be particularly high in the warm season. Research into metabolic dependency of bio-mediated sediment resuspension may help to place phenomenological observations in the broader framework of metabolic theories in ecology and to formulate general expectations on the coastal ecosystem functioning.

Restoration of biogeomorphic systems by creating windows of opportunity to support natural establishment processes.

In degraded landscapes, recolonization by pioneer vegetation is often halted by the presence of persistent environmental stress. When natural expansion does occur, it is commonly due to the momentary alleviation of a key environmental variable previously limiting new growth. Thus, studying the circumstances in which expansion occurs can inspire new restoration techniques, wherein vegetation establishment is provoked by emulating natural events through artificial means. Using the salt-marsh pioneer zone on tidal flats as a biogeomorphic model system, we explore how locally raised sediment bed forms, which are the result of natural (bio)geomorphic processes, enhance seedling establishment in an observational study. We then conduct a manipulative experiment designed to emulate these facilitative conditions in order to enable establishment on an uncolonized tidal flat. Here, we attempt to generate raised growth-promoting sediment bed forms using porous artificial structures. Flume experiments demonstrate how these structures produce a sheltered hydrodynamic environment in which suspended sediment and seeds preferentially settle. The application of these structures in the field led to the formation of stable, raised sediment platforms and the spontaneous recruitment of salt-marsh pioneers in the following growing season. These recruits were composed primarily of the annual pioneering Salicornia genus, with densities of up to 140 individuals/m2 within the structures, a 60-fold increase over ambient densities. Lower abundances of five other perennial species were found within structures that did not appear elsewhere in the pioneer zone. Furthermore, recruits grew to be on average three times greater in mass inside of the structures than in the neighboring ambient environment. The success of this restoration design may be attributed to the combination of three factors: (1) enhanced seed retention, (2) suppressed mortality, and (3) accelerated growth rates on the elevated surfaces generated by the artificial structures. We argue that restoration approaches similar to the one shown here, wherein the conditions for natural establishment are actively mimicked to promote vegetation development, may serve as promising tools in many biogeomorphic ecosystems, ranging from coastal to arid ecosystems.

Management of local stressors can improve the resilience of marine canopy algae to global stressors.

Coastal systems are increasingly threatened by multiple local anthropogenic and global climatic stressors. With the difficulties in remediating global stressors, management requires alternative approaches that focus on local scales. We used manipulative experiments to test whether reducing local stressors (sediment load and nutrient concentrations) can improve the resilience of foundation species (canopy algae along temperate rocky coastlines) to future projected global climate stressors (high wave exposure, increasing sea surface temperature), which are less amenable to management actions. We focused on Fucoids (Cystoseira barbata) along the north-western Adriatic coast in the Mediterranean Sea because of their ecological relevance, sensitivity to a variety of human impacts, and declared conservation priority. At current levels of sediment and nutrients, C. barbata showed negative responses to the simulated future scenarios of high wave exposure and increased sea surface temperature. However, reducing the sediment load increased the survival of C. barbata recruits by 90.24% at high wave exposure while reducing nutrient concentrations resulted in a 20.14% increase in the survival and enhanced the growth of recruited juveniles at high temperature. We conclude that improving water quality by reducing nutrient concentrations, and particularly the sediment load, would significantly increase the resilience of C. barbata populations to projected increases in climate stressors. Developing and applying appropriate targets for specific local anthropogenic stressors could be an effective management action to halt the severe and ongoing loss of key marine habitats.