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Vegetation Establishment Following Floodplain Restoration in Mediterranean-climate California

Abstract

Although herbaceous communities are important components of floodplain ecosystems, the factors constraining their restoration and post-restoration dynamics are poorly understood. Over the decade following restoration of a 3.2 km reach of the Merced River and floodplain in California, we tracked herbaceous community composition to distinguish floodplain habitats and utilized perturbations from revegetation treatments and post-restoration flooding to generate community assembly rule hypotheses regarding treatment effectiveness and persistence, with a particular interest in native perennials capable of suppressing non-natives over time if undisturbed. Revegetation treatments comprised combinations of sowing a sterile cover crop, sowing native species, and inoculating mycorrhizae. Most surveyed floodplain areas comprised a low terrace characterized by exceptionally droughty soils, relatively deep groundwater, and occasional flooding lasting into summer. Few species could tolerate both flood and drought to this extent, and the flood year community was generally distinct from that in non-flood years. Both communities were dominated by ruderals capable of avoiding stress and re-establishing following disturbance, including many non-native annual grassland species. Only Artemisia douglasiana responded to the treatments, as most seeded native species failed to establish, including those native perennial grasses expected to suppress non-native annuals, while other seeded native species either established adequately from natural dispersal or failed to persist through moderate flooding. Neither the cover crop nor mycorrhizal inoculation had any meaningful effect. Restoration efforts in naturally ruderal-dominated habitats may be better spent allowing natural regeneration, addressing particularly noxious invasives, and identifying or constructing habitats supporting long-lived native perennials.

Although originally developed for population sizes and population growth rates, modern capture-recapture models can estimate demographic rates in complex situations: multistate models for multiple study sites and stage-structured populations, superpopulation entry probability models for recruitment, and multievent models when state assessments are uncertain. However, combinations of these complications, such as recruitment studies with uncertain state assessments, are common, yet no single model has explicitly incorporated all of these elements. Ultimately, these models estimate the same fundamental population process with the same general approach, and we combine them in a generalized hidden process model based upon a simple discrete state and transition population model with Poisson recruitment that can estimate how recruitment and survivorship rates vary with respect to measured covariates from uncertain state assessments for a stage-structured population at multiple sites. Although closely related to the motivating models, the generalized model relaxes the Markov assumption. While we provide the distributions necessary to implement Bayesian data augmentation methods, we also provide an efficient analytical likelihood with a compact parameter space that is applicable in the absence of density-dependent mortality. As a demonstration, we estimate the influence of several covariates on recruitment and survivorship rates from uncertain observations of Salix gooddingii seedlings at different locations along a riparian gradient, and we use simulations to examine variation in the precision of estimated parameters.

In Mediterranean climates, cottonwoods and willows often exhibit high germination and seedling mortality rates, with recruitment occurring primarily in the occasional year when favorable spring floods improve survivorship. However, along the Robinson Reach of the Merced River, both germination and mortality rates appeared to be atypically low. To understand why these rates were so low along this recently restored flow-regulated, gravel-bedded stream, we surveyed Populus fremontii, Salix exigua, and Salix gooddingii, estimated germination and survivorship rates, and examined their correlations with factors expected to constrain recruitment, namely seed release, seed arrival, moist germination beds, light levels, groundwater depth, groundwater recession rates, and shear stress. Germination/initial establishment rates were low due in part to low seed arrival rates. Only Salix gooddingii was abundant enough to model in detail, and while moist germination surfaces increased germination/initial establishment, rates were low overall. Survivorship rates for Salix gooddingii seedlings and for small individuals were not correlated with any examined covariates. Seedlings tolerated moderate competition, and the absence of major scouring, even during 6 year flows, enabled survival at sites with sufficiently shallow groundwater that seedlings were unaffected by groundwater recession rates.

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