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The Role of Seed Bank and Germination Dynamics in the Restoration of a Tidal Freshwater Marsh in the Sacramento–San Joaquin Delta Taylor M

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The Role of Seed Bank and Germination Dynamics in the Restoration of a Tidal Freshwater Marsh in the Sacramento–San Joaquin Delta Taylor M SEPTEMBER 2019 Sponsored by the Delta Science Program and the UC Davis Muir Institute RESEARCH The Role of Seed Bank and Germination Dynamics in the Restoration of a Tidal Freshwater Marsh in the Sacramento–San Joaquin Delta Taylor M. Sloey,1,2 Mark W. Hester2 ABSTRACT enhanced using a pre-germination cold exposure, Liberty Island, California, is a historical particularly for species of concern for wetland freshwater tidal wetland that was converted restoration. The cold stratification study showed to agricultural fields in the early 1900s. that longer durations of pre-germination cold Liberty Island functioned as farmland until an enhanced germination in Schoenoplectus acutus, accidental levee break flooded the area in 1997, but reduced germination in Schoenoplectus inadvertently restoring tidal marsh hydrology. californicus, and had no effect on Typha latifolia. Since then, wetland vegetation has naturally Overall, germination of S. californicus and recolonized part of the site. We conducted a seed S. acutus was much lower than T. latifolia. Our bank assay at the site and found that despite a findings suggest that seeding may not be an lack of germination or seedling recruitment at effective restoration technique for Schoenoplectus the site, the seed bank contained a diverse plant spp., and, to improve restoration techniques, community, indicating that the site’s continuous further study is needed to more comprehensively flooding was likely suppressing germination. understand the reproduction ecology of important Additionally, the frequency of germinating seeds marsh species. in the seed bank did not represent the dominant adult plant community. We conducted a cold stratification study to determine if this observed KEY WORDS disparity could be explained by seed germination cold stratification, dormancy, restoration, dynamics, and whether germination could be Schoenoplectus, self-design, tidal freshwater, Typha SFEWS Volume 17 | Issue 3 | Article 5 INTRODUCTION https://doi.org/10.15447/sfews.2019v17iss3art5 Worldwide, tidal freshwater wetlands have been in * Corresponding author: [email protected] decline as a result of human alteration of natural 1 Division of Science hydrological regimes (Zonneveld 1960; Dahl 1990; Yale-NUS College Singapore 138533 Kotze et al. 1995; An et al. 2007; Barendregt et 2 Department of Biology al. 2008; Davidson 2014). Efficient restoration University of Louisiana, Lafayette of these degraded wetlands and creation of new Lafayette, LA, USA 70504 wetlands habitats is desired globally to restore the important ecosystem functions these systems SAN FRANCISCO ESTUARY & WATERSHED SCIENCE VOLUME 17, ISSUE 3, ARTICLE 5 provide. However, many candidate wetland Seed banks represent a composite of seeds of restoration sites have undergone severe alteration varying degrees of dormancy, and play an to their physical and biological attributes, important role in the long-term and short-term inhibiting their ability to achieve restoration preservation of species at a location (Baskin goals. The need for successful restoration of tidal and Baskin 1978; Bigwood and Inouye 1988; wetlands reflects a larger demand to rehabilitate van der Valk and Rosburg 1997; Peterson and and restore degraded ecosystems in general. The Baldwin 2004). Because of limitations on seed United Nations General Assembly announced dispersal and germination, the seed bank of a earlier this year that 2021–2030 will be the site may include a greater diversity of species Decade on Ecosystem Restoration (UN 2019). than what is actually present in the standing However, knowledge gaps still exist in basic adult plant community (Hopfensperger 2007). plant ecology, which will be crucial to address, Understanding the species in the seed bank — and to ensure successful science-based ecosystem limitations to seed establishment — is crucial for restoration, particularly in wetlands. wetland restoration or creation projects (van der Valk et al. 1992). Both self-design and designer In tidal freshwater wetlands, three factors are restoration strategies would benefit from a better generally understood to be the most influential understanding of seed ecology and germination in controlling ecosystem structure and function: dynamics of key plant species as a means to hydrology, species dispersal, and biological improve seeding/planting approaches (Zedler dynamics of the species (i.e., ability to cope with 2000) or establish the proper abiotic conditions to abiotic conditions) (Grubb 1977; Harper 1977; van facilitate natural recruitment (Palmer et al. 1997). der Valk 1981). Site elevation and corresponding flooding characteristics are often the most Seeding, or the intentional introduction of seeds influential physical parameters that control the of a target species, is a common restoration establishment of the plant community (Barendregt strategy in wetlands (Mitsch et al. 1998; Ruiz- et al. 2008; Mitsch and Gosselink 2015). Jaen and Aide 2005). Because of dormancy and Re-creating the proper hydrology and physical germination dynamics in desired species, the use parameters is an essential first step to wetland of pre-germination seed treatments — such as restoration, but it is only part of a more complex cold-stratification, dark conditioning, physical combination of factors that must be addressed scarification, bleach scarification, or repeated (Hilderbrand et al. 2005). Seed dispersal, cycles of imbibition — may be implemented germination, and seed abundance can further to accelerate germination or increase total limit restoration outcomes after re-introduction percent germination of seeds. These techniques of wetland hydrology (van Leeuwen et al. 2014). have been used on a wide variety of species, Meta-analyses compiled from a variety of including agriculturally important crops and ecosystems suggest that colonization of primary target restoration species, throughout diverse successional species depends on the interplay biogeographical ranges (Heydecker et al. 1973; between the propagule and abiotic and biotic Schütz and Rave 1999; Kaye and Kuykendall limitations on seedling establishment (Seabloom 2001; Kulkarni et al. 2006). However, data gaps 2011; Titus and Bishop 2014). Additionally, still remain regarding the seed ecology and abiotic conditions may differentially limit plant germination dynamics of species important to recruitment, depending on species and life- freshwater tidal marshes. history stage (Shumway and Bertness 1992; Bunn and Arthington 2002; Hinkle and Mitsch In this study, we implemented a seed bank assay 2005). To understand community structure and at Liberty Island, a post-levee breach wetland restoration dynamics, propagule limitations and restoration site in the Sacramento San-Joaquin abiotic constraints on plant recruitment must be Delta in California. To complement our seed bank understood (Moore and Elmendorf 2006). assay, we also conducted a cold stratification study to understand germination dynamics and determine whether germination in three tidal 2 SEPTEMBER 2019 marsh plant species could be artificially improved characteristic of California’s Mediterranean with this technique. Additionally, to understand climate, the intention of our study was to how hydrology may limit seed germination, we determine if pre-germination treatments could be conducted an inundation study in which we used to artificially improve species germination; exposed seeds (that had already been exposed restoration practitioners want this to improve the to various pre-germination cold stratification practices of restoration by seeding. Furthermore, treatments) to varying depths of flooding. understanding how cold periods affect the way species respond to seed germination may inform Our controlled germination studies used seeds understanding of species regeneration and from three emergent tidal marsh macrophyte colonization dynamics at broader regional scales. plant species that are native to California and The objective of our study was to (1) determine commonly implemented in restoration plantings: if viable wetland seeds existed in the seed bank California bulrush (Schoenoplectus californicus, of the Liberty Island mudflats, (2) determine if family: Cyperaceae), hardstem bulrush cold treatment could enhance germination for (Schoenoplectus acutus, family: Cyperaceae) three dominant wetland plant species, and (3) and broadleaf cattail (Typha latifolia, family: understand how hydrology affects germination Typhaceae). All three of these species are in those three wetland plant species. The classified as obligate wetland plant species (USDA overarching goal of these combined studies was 2019) and tolerate being inundated > 60% of the to increase understanding of seed-germination day (Sloey et al. 2016). However, much less is requirements and colonization dynamics in known about the hydrologic conditions needed tidally-influenced wetlands to maximize for germination and seedling recruitment in these restoration efficiency in similar systems in the species. Studies in other regions have reported future. high seed germination rates in Typha latifolia (Sifton 1959; Keddy and Ellis 1985), whereas few studies have examined Schoenoplectus spp. MATERIALS AND METHODS germination dynamics. Kellogg et al. (2003) Seed Bank Assay found that S. acutus seeds did not germinate Site Description when flooded, and showed very low percent Liberty Island is a freshwater (salinity < 0.2 ppt) germination (1%) even when not inundated. tidal marsh restoration site in the
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