Change in fish functional diversity and assembly rules in the course of tidal marsh restoration

Functional trait theory provides a mechanistic framework to understand change in community composition and community assembly through time and space. Despite this, trait-based approaches have seldom been used in ecological restoration. Succession theory predicts that habitat complexity and resource availability will increase with restoration time, leading to increased functional dissimilarity among coexisting species. However, in the case of tidal marsh restoration, it is not clear whether reestablishing the harsh abiotic conditions typical of estuaries will initiate successional trajectories. We investigated monotonic changes in the functional structure of fish communities and shifts in assembly mechanisms, with tidal restoration time. A five-level gradient of 'intertidal habitat naturalness' was constructed from a set of artificialized (dyked), restored (with different ages) and natural intertidal sites, and used as a surrogate for restoration progress. The fish ecophases were described using ten functional traits related to food acquisition and swimming ability. The trends in six functional dimensions (identity, richness, evenness, dispersion, originality and specialization) were investigated along the naturalness gradient. Consistenly with succession theory, functional specialization, dispersion and, less markedly, richness increased with intertidal naturalness meaning that restored and natural intertidal habitats supplied fish with specific foraging and dwelling conditions absent from dyked marshes. Community assembly patterns varied with respect to traits and differed at both ends of the naturalness gradient. Dyked marshes were more affected by trait convergence possibly due to limiting resources. Environmental filtering was detected all along the naturalness gradient although the traits affected varied depending on the naturalness level of habitats. Environmental filtering tended to decrease in restored and natural intertidal habitats. Increased naturalness restored the attractivity of benthic habitats as feeding or settling grounds, promoted shelter-seeking vs. free-swimming strategists and favoured ecophases with carnivorous diets, feeding on microinvertebrates and benthic low-mobility macroinvertebrates. Approaches based on functional trait diversity have the potential to question and refine the theoretical frame of ecological restoration and to assist managers in their efforts to restore tidal wetlands.