Active Versus Passive Revegetation on Recently Restored Tidal Marsh Hummocks: Does Planting Make a Difference?

By: Corinne Kennah, Katie White, Zachary Silber-Coats

Abstract
The Wood Creek property in Humboldt County, California is the site of a recent tidal marsh restoration project. The property is owned by the Northcoast Regional Land Trust (NRLT), and in 2010 the land trust conducted a restoration project that reestablished historic tidal influence to the Wood Creek property, which had been used as pastureland for decades. This study compared and analyzed the effectiveness of two revegetation techniques on eight constructed tidal marsh hummocks, six of which were actively seeded and planted with native species while the remaining two were left as-to to allow vegetation to passively establish over time. We found that three years after the restoration there was no significant difference in total colonization by exotic vegetation on actively seeded and planted hummocks compared to hummocks that were allowed to passively revegetate over time. The dominant species on both hummock types was the nonnative pasture grass, Agrostis stolonifera. Planted and seeded native varieties did not flourish. Some success of active revegetation efforts were seen by the decreased presence of A. stolonifera on actively revegetated hummocks in relation to passively revegetated hummocks, and also by a greater abundance of Deschampsia cespitosa subs. cespitosa (an actively planted and seeded native species) on actively revegetated versus passively revegetated hummocks. However, although these differences were visible in the field as well as in our data, analysis did not find that these differences were significant.

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Introduction
According to the U.S. Fish and Wildlife Service, wetlands can be characterized by a water regime intermediate between those of terrestrial and aquatic systems (Zedler and Kercher, 2005). Estuarine systems are characterized as aquatic systems located at the interface between freshwater and oceanic environments (Schlosser and Eicher, 2012). Brackish and salt marsh habitats, as estuarine wetlands, thus form valuable edge habitats with respect to both salinity and hydric influence. Coastal wetland habitats are crucial to global biodiversity (Zedler and Kercher, 2005). Important ecological services of coastal wetlands include water purification via nutrient uptake, amelioration of the effects of severe storms and floods, and groundwater recharge (Zedler and Kercher, 2005). Prominent biodiversity services provided by coastal wetlands include habitat for juvenile fish and invertebrates of economic importance, alongside the crucial role that these habitats play in both breeding and wintering habitat for migratory waterfowl (Zedler and Kercher, 2005). It is estimated that approximately half of all wetlands present worldwide have been lost to competing land uses since the 1800s, and coastal wetland loss may be greater (Zedler and Kercher, 2005). The current extent of salt marsh ecosystems in North America is only a small fragment of what occurred historically. Loss and degradation has occurred since European settlement from land use changes and disturbances such as farming, grazing, dredging, and altering of natural hydrologic regimes and sediment supply (Gedan et al., 2009). It has been conservatively estimated that 50% of all historic salt marshes in the United States have already been lost (Kennish, 2001). Laws enacted in recent decades by federal and state governments

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have helped curtail large scale impacts to the nation's wetlands through direct causes. However, non-direct impacts from exotic species invasion, eutrophication and climate change related sea level rise are now beginning to be seen as major sources of salt marsh loss and degradation (Gedan et al., 2009). In Humboldt Bay, salt marsh loss has been particularly dramatic. Since the beginning of European settlement in the area in the 1850s, over 90% of Humboldt Bays salt marsh has been diked, drained, and converted to pasture land or developed (Pickart, 2001). Prior to European settlement, Humboldt Bay had 9,000 acres of salt marsh habitat (Pickart, 2001). Following settlement, the railroad, and extensive agricultural land conversions, Humboldt Bay now has less than 900 acres of salt marsh remaining (Pickart, 2001). Salt marsh in Humboldt Bay has not only faced direct destruction due to lack of tidal influence, but is also threatened by a highly invasive species. Dense-flowered cordgrass (Spartina densiflora) arrived in Humboldt Bay in the late 1800s, most likely from ship ballast carried from Chile (USFWS, 2013). As of 2001, S. densiflora occupied 94% of the 900 acres of remaining Humboldt Bay salt marsh (Pickart, 2001). It was first identified as an invasive species in 1984 (USFWS, 2013). Invasive species such as S. densiflora drastically outcompete native vegetation and therefore limit salt marsh plant diversity (USFWS, 2013). This is of particular concern in the high salt marsh habitats of Humboldt County, which support over 20 native plant species (USFWS, 2013).The high level of salt marsh degradation around Humboldt Bay highlights the need for restoration. Restoration projects in the area tend to focus either on converting agricultural land back to salt marsh, or on removing invasive species from existing salt marsh.

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Post-project monitoring is an important aspect of every ecological restoration project. Monitoring allows for evaluation of the project in order to assess impacts and the degree to which restoration goals have been met. Post-project monitoring can also help identify areas that require further treatment or a different management approach (Bash and Ryan, 2002; Thom, 2000). Post-project monitoring also has overarching importance in that the reporting of results can help drive the success of future restoration projects and restoration knowledge (McTigue et al., 2005) Aquatic restoration projects are especially susceptible to failure (Thom, 2000). For this reason, monitoring of projects in these ecosystems is of utmost importance. Unfortunately, monitoring efforts on restoration projects are often constrained by a lack of funding. However, large sums of money are now being spent on restoration activities. Between 2000 and 2007, over $100 million dollars was allocated to restoration efforts in Humboldt County (Baker and QuinnDavidson, 2011). With so much being spent on restoration annually, it is important to increase the success of projects so that restoration efforts are not done in vain. The objectives of this study were to examine vegetation on tidal hummocks in the Wood Creek Tidal Marsh Enhancement Project. The main objective was to compare current vegetation on hummocks that were actively revegetated versus vegetation that passively colonized on hummocks that were not actively revegetated.

Methods
Site Description and Characteristics
The Wood Creek Tidal Enhancement project site is located north of Myrtle Avenue, northeast of Eureka, California (Figure 1). The project is located on over 35 acres on the western portion of the Wood Creek property (McBain and Trush, 2007).

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Figure 1. Map of the Wood Creek property, showing the project site and constructed slough channels.

In 2005, the Northcoast Regional Land Trust (NRLT) acquired 54 acres of former tidal salt marsh (NRLT, 2012). In the late 1800s or early 1900s, the land was diked by European settlers in order to turn the salt marsh into viable agricultural land for growing crops and grazing livestock (Schlosser and Eicher, 2012). The land has been farmed in the past, and in more recent decades has been a cattle pasture (ODowd A; personal communication, 2013). Prior to site purchase in 2005 by the NRLT, the site was owned by Freshwater Farms. It was used to grow many different species of native trees to provide propagules for sale at Freshwater Farms Nursery, a local

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restoration nursery that also provided thousands of propagules for the brackish marsh enhancement. The climate of the site is transitional between cool-summer Mediterranean and moist maritime zones (McBain and Trush, 2007). Temperature variation is remarkably small; summer maximum temperatures rarely exceed 70F, and winter minima rarely drop below 30F (NWSFO-NOAA, 2013). Rainfall is approximately 40 inches per year, concentrated between October and April (NWSFO-NOAA, 2013). Soil at the project site is a silty clay loam of the Occidental series (USDA-NRCS, 2013). Depth to the aquifer is commonly one meter or less, and surface water is frequently present (USDANCCS, 2013). Current site elevation ranges between approximately 6.0 to 7.5 feet above mean lower low water (MLLW), placing the site towards the upper limit of tidal influence (McBain and Trush, 2007).

Overview of Study Methods

For this study we assessed post-restoration vegetation on four tidal hummocks. Two of the hummocks were actively revegetated after restoration, while the other two hummocks were not revegetated, and were instead left as controls to observe the passive colonization of vegetation naturally over time. Data for actively revegetated hummocks was then compared with data from passively revegetated hummocks to determine whether initial planting methods had a significant influence on hummock vegetation. This study was conducted three years after final restoration actions were implemented, leaving a substantial amount of time for hummock vegetation to develop.

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Restoration Project Background

The Wood Creek Estuary Revegetation Project was a subset of a larger project, the Wood Creek Tidal Marsh Enhancement Project. According to the Wood Creek Tidal Marsh Design Report for the Wood Creek Tidal Marsh Enhancement Project (Anderson and Associates, 2008), the goals of this project were to reestablish historic tidal hydrology to the site by opening the tide gate located at the confluence of Wood Creek and Freshwater Slough and reestablishing a network of slough channels (Figure 2). Through these actions, the project area was subject to tidal influence for the first time in almost a century. Improvement of winter rearing habitat for salmonids, such as Coho salmon (Oncorhynchus kisutch) and steelhead trout (Oncorhynchus mykiss), was also a central goal of the restoration project (NRLT, 2012). Winter juvenile rearing habitat was improved by increasing the cover of native wetland plant species, and increasing access to areas of refuge from high velocity flow events by establishing connections to floodplains, side channels, and off-channel ponds. Off-channel ponds were also created with the objective of increasing and improving habitat for the federally endangered tidewater goby (Eucyclogobius newberryi) that has been seen on the site (NRLT, 2012). Heavy equipment was used to excavate slough channels, create off channel ponds, remove a 300 foot berm, and build eight tidal hummocks in the wetland area using sediment excavated throughout the project site, reducing the exepenses associated with trucking sediments off-site (NRLT, 2012). Six of the eight hummocks were actively revegetated with seed and plugs grown at Freshwater Farms Nursery, while the remaining two hummocks were left as-is to allow for the monitoring of passively colonized vegetation over time.

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Figure 2. Map showing restoration design elements of the Wood Creek Tidal Marsh Enhancement Project, the larger project of which the Wood Creek Estuary Revegetation Project was a subset to (Image source: Anderson and Associates, 2008). Actively revegetated hummocks were seeded and planted with a few different native brackish habitat plant species (NRLT, 2012). The NRLT planted 37,000 plants on the actively revegetated hummocks, including Deschampsia caespitosa (tufted hairgrass), Carex lyngbyei (Lyngbyes sedge), silverweed Potentilla anserina (silverweed), Distichlis spicata (saltgrass), Juncus effusus (softstem rush), and Scirpus microcarpus (small-fruited bulrush) (NRLT, 2012). It was difficult to find post-project information on the specific modes of planting ultimately used, but a biological assessment completed in 2007 reported the NRLTs original plans (McBain and Trush, 2007). According to the report, the NRLT intended to plant seeds and nursery stock of Deschampsia cespitosa ssp. cespitosa (tufted hairgrass), Carex lyngbyei (Lyngbye sedge), and nursery stock of Potentilla anserina (silverweed), Distichlis spicata (saltgrass), and Juncus
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lescurii (salt rush; formerly J. lesueurii) (McBain and Trush, 2007). It is unclear whether or not these species were ultimately planted in the modes specified by the 2007 assessment, but this information may indicate the approximate forms of outplanting propagules.

Field Methods
We conducted line transects to compare vegetation on actively vegetated versus passively vegetated hummocks within the restoration site. To do this, we recorded vegetation data on two passively vegetated hummocks and two actively vegetated hummocks, as indicated in Figure 3. On each hummock, we conducted at least two transects, the combined length of which totaled 200 feet per hummock (Figure 4). The number of transects and individual transect lengths on each hummock was determined by the size and shape of each hummock. The transect locations were determined by a stratified sampling method (Neyman, 1934). Once the first transect on a hummock was placed, we then placed all following transects parallel to the first at a distance of 20 feet apart. The location of the first transect on each hummock was determined by first defining the border of the hummock and then measuring 30 feet from the northwestern edge of the hummock, to eliminate edge effects. Dominant and subdominant vegetation were identified directly under the transect tape until we reached a distinct change in dominant vegetation. Each change in dominant vegetation was considered a separate sampling unit within each transect. We then calculated percent cover of dominant and subdominant vegetation on each hummock by comparing the total length occupied by each species from each transect to the total length of all the transects (200 feet). We then used these data to analyze the success of native vegetation versus nonnative or invasive vegetation on the two different types of hummocks. To analyze our data, we ran t-tests of log-normalized percent cover by species and made bar graphs to compare

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the percent cover of each dominant or subdominant species between actively revegetated and passively revegetated hummocks.

Figure 3. Map showing the tidal hummocks that were constructed within the project area, distinguishing between actively revegetated and passively revegetated hummocks. The map also shows the four hummocks that were surveyed for this study (Image source: NRLT, 2009).

Figure 4. Map showing approximate locations of transects conducted on each hummock surveyed (Image source: NRLT, 2009).

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Results
Surveyed transects on hummocks included both native and non-native plant species. Plant species found, and their status as native or non-native, are listed below (Table 1). Table 1. Plant species present on transects surveyed.
Latin name Agrostis stolonifera Holcus lanatus Lotus corniculatus Elymus repens Rumex sp. Agrostis sp. Hordeum brachyantherum Atriplex sp. Juncus lesueurii Carex lyngbyei Aster chilensis Triglochin maritima Deschampsia cespitosa ssp. cespitosa Common name Creeping bentgrass Velvet grass Birdsfoot trefoil Quackgrass Dock Bentgrass Native barley Salt bush Salt rush Lynbye Sedge Chilean Aster Arrow-grass Tufted hairgrass Native status Non-native Non-native Non-native Non-native Non-native Non-native Native Native Native Native Native Native Native

Non-native Agrostis stolonifera (creeping bentgrass), an exotic rangeland grass, was the most common plant species observed on both actively revegetated and passively-revegetated transects, forming approximately 40% and 80% of the total transect distances, respectively. Native Deschampsia cespitosa subsp. cespitosa (tufted hairgrass) was the second most common species, forming over 15% of surveyed vegetation on passively revegetated hummocks, and approximately 30% on actively revegetated hummocks. Non-native Holcus lanatus (velvet grass) was a substantial component of actively revegetated hummocks, but nearly absent from passively

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revegetated hummocks. All other species occurred sporadically and as small percentages of the total (Figure 5). Agrostis stolonifera was prominent as a dominant component of both actively and passively revegetated hummocks. However, A. stolonifera occupied nearly twice as much of the passively revegetated hummocks, as compared to its presence in actively-revegetated hummocks. Conversely, Holcus lanatus, also an invasive grass, was dominant on over a quarter of the linear feet surveyed in actively revegetated hummocks, but all but absent on passively revegetated hummocks.

Figure 5. Dominant nonnative (exotic) and native plant species by percentage of total transect length, in actively and passively restored hummocks.
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Agrostis stolonifera was also prominent as a subdominant component of vegetation on both hummock types (Figure 6). In general, subdominant vegetation along transects was more broadly divided among species than was the dominant vegetation, in the case of both actively and passively revegetated hummocks.

Figure 6. Subdominant exotic and native plant species by percentage of total transect length, in actively and passively restored hummocks. We conducted paired t-tests of log-normalized vegetation percentages for the three most common plant species to evaluate the null hypothesis that mean measured length of dominant

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plant species did not differ between actively and passively revegetated hummocks. Results of this test were insignificant, suggesting that transects on active and passive hummocks did not differ conclusively in the three most common dominant plant species (t = 0.9319, df = 2.019, p = 0.449). Results of a similar t-test on subdominant vegetation were similarly insignificant (t=1.1626, df = 2.869, p = 0.3325). We then calculated a Spearmans rank-correlation coefficient (rho) to determine the extent to which percentages of plant species on passively revegetated hummocks correlate with species percentages on actively-revegetated hummocks. Spearman coefficients for exotic and native species are reported below (Table 2). Table 2. Spearmans correlation coefficient (Rho) for subdivisions of plants surveyed.
Plant type Non-native Non-native Native Native Vegetation Component Dominant Subdominant Dominant Subdominant Spearmans Rho 0.169 0.514 0.368 0.419

The low Spearman correlation coefficient for exotic dominant plant species indicates that dominant exotic plant composition differs between actively and passively revegetated hummocks. Higher Spearman correlation coefficients for dominant native vegetation, and for subdominant vegetation both native and exotic, indicate that vegetation is more similar by hummock type for these vegetation categories, as compared to the dominant exotic vegetation.

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Discussion
Before delving into the potential implications of our findings, there are a few factors which are important to take into consideration. Our transect results may have been affected by the seasonal timing of our survey. We observed large areas of dead thatch underneath much of the vegetation we surveyed. While this could in many cases have belonged to the same species as the actively growing vegetation, much of this thatch was in an advanced state of decomposition, rendering moot any attempts at identification. In addition, we observed evidence that several species in the larger marsh habitat had already fully senesced. In particular, dead leaves of dormant Potentilla anserina were conspicuous in several areas outside the hummocks. Of the native species sown and planted on the actively revegetated hummocks, only D. cespitosa ssp. cespitosa was particularly successful. At 30% dominance, D. cespitosa ssp. cespitosa had twice the percent dominance on actively revegetated hummocks, as compared to 15 % on passively revegetated hummocks. Neither C. lyngbyei nor J. lescurii achieved a dominant presence on actively revegetated hummocks, though they were 5% and 4% dominant on passively revegetated hummocks, respectively. While the planting of D. cespitosa ssp. cespitosa appears to have had an impact on the species success, planting of other native species appears to have had no impact. The lack of success among native species other than D. cespitosa ssp. cespitosa on actively revegetated hummocks suggests that the revegetation strategy employed could have been more economically efficient. Specifically, restoration practitioners might have had more success by planting native species better able to compete with invasive grasses. Based on the results of coastal prairie plots in more southerly areas of California, inclusion of fastergrowing native prairie plants such as Calamagrostis nutkaensis (Pacific reedgrass), and Festuca rubra (red fescue), would likely have improved exclusion of invasive grass species (Corbin et

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al., 2004). Differences in seed germination, growth, and seed bank may each have had a role in the lack of success of the planted native species at this site. The low success rate of native seeds and plugs may also be due to the significant presence of the invasive species, A. stolonifera. It is likely that much of the native seed and plants were outcompeted by A. stolonifera, which was found to have the highest percent cover of all species on both actively and passively revegetated hummocks (80% and 40% respectively). Dominant invasive vegetation between actively and passively restored hummocks was found to have a lower Spearman correlation coefficient (0.169) than other components of vegetation sampled, suggesting high dissimilarity in invasive plant cover between hummock types. This can be explained primarily by the predominance of H. lanatus on approximately 25% of the transect length in actively restored hummocks. In contrast, this species was nearly absent on passively revegetated hummocks. If we consider the sum of A. stolonifera and H. lanatus percentages on actively revegetated hummocks, invasive grasses occupied 65% of transect length on these hummocks, as compared to 80% of transect length on passively restored hummocks. According to the results of the t-test of dominant vegetation, there is not a significant difference in total cover of these two invasive species between hummock types, so we can interpret the approximately 15% difference in invasive grass cover as insignificant. The dominant presence of A. stolonifera on the hummocks is not surprising considering the nature of the species. Agrostis stolonifera is a perennial pasture grass that reproduces both sexually by seed and vegetatively through stolons (Zapioloa et al., 2008). Unlike many perennial grasses, A. stolonifera is able to set seed in a single growing season (Esser, 1994). This allows for the establishment of a large bank. In addition to having a large seed bank, A. stolonifera seeds

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are very small, making them more easily dispersed by wind than many other grasses (Zapiola et al., 2008). Due to the ease of dispersal through both seed and stolons, A. stolonifera readily colonizes disturbed sites. The species is also known to exhibit high levels of phenotypic plasticity (Zapiola et at., 2008), making it extremely tolerant to changing environmental conditions and management actions employed to control the spread of the species. The seedbank of A. stolonifera is viable in soil for one to four years (Esser, 1994; Zapiola et al., 2008). The material used to create the eight tidal hummocks on the project site came from the sediment excavated during restoration activities. The site was formerly used as a cattle pasture for several decades prior to the restoration project (ODowd, A.; personal communication, 2013), and because of this, the sediment was probably dominated by the seed banks of pasture grasses such as A. stolonifera and H. lanatus. Therefore, it is likely that a viable A. stolonifera seedbank existed in the excavated soils that were used to build the hummocks. Since D. cespitosa spp. cespitosa was dominant on substantial proportions of both hummock types, its success may also be due to the existing seedbank for this species. Despite the higher success rate of D. cespitosa spp. cespitosa on actively revegetated hummocks than on passively revegetated hummocks, passively revegetated hummock transects suggest that the species can compete with reasonable success against A. stolonifera up to a 15% cover dominance without the need for active outplanting. Although revegetation efforts may have decreased the success rate of A. stolonifera, the apparently lower success rate of A. stolonifera on actively revegetated hummocks than on passively revegetated hummocks might be attributed to the relative success of another invasive grass, Holcus lanatus, or might instead reflect the relatively greater success of native

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Deschampsia cespitosa ssp cespitosa. While the active revegetation was not successful in repopulating the hummocks with only native species, the slight difference in A. stolonifera success suggests that further research and management actions might yield effective strategies for limiting A. stolonifera on the site. The successful predominance of Agrostis stolonifera on both hummock types, and the low percent dominance of planted and unplanted native species, indicate that the active revegetation conducted by the NRLT was somewhat ineffective. However, as suggested in a grant proposal prepared by NRLT for the project, the revegetation effort can still benefit restoration knowledge as a whole. By collecting data on the success of the different hummock types, the effort will provide background research on the effectiveness of active revegetation on excavated soils, particularly in historical brackish marshland impacted by former agricultural use. Future research may lead to insight on effective strategies to decrease A. stolonifera success, to mitigate negative impacts of revegetation (such as the increased success of H. lanatus on actively revegetated hummocks), in addition to insight as to outplanting choices better able to establish in favor of invasives. Long-term monitoring at the site may also yield results in contrast to earlier data. For instance, it is possible that planted native seed or plugs may result in the establishment of those native species several years after this study. Given the findings of this study, similar restoration projects conducted in the future may save resources by either foregoing active restoration or researching and utilizing native plant species known to be competitive with invasive pasture grasses, such as Calamagrostis nutkaensis (Pacific reedgrass), and Festuca rubra (red fescue) in their active revegetation efforts (Corbin et al., 2004).

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Northcoast Regional Land Trust (NRLT). 2009. Wood Creek Estuary Revegetation Project. Grant proposal for TNC-NOAA Community-Based Habitat Restoration Grants. Pickart, Andrea. 2001. The distribution of Spartina densiflora and two rare salt marsh plants in Humboldt Bay 1998-1999. U.S. Fish and Wildlife Service, Humboldt Bay National Wildlife Refuge. Schlosser, S. and A. Eicher. 2012. The Humboldt Bay and Eel River Estuary Benthic Habitat Project. California Sea Grant Publication T-075. <http://wwwcsgc.ucsd.edu/bookstore/documents/HumboldtLR.pdf> Accessed October 22, 2013. Thom, R.M. 2000. Adaptive management of coastal ecosystem restoration projects. Ecological Engineering 15: 365-372. United States Department of Agriculture, National Cooperative Soil Survey (USDA-NCCS). 2013. Official series description: Occidental. <https://soilseries.sc.egov.usda.gov> Accessed October 30, 2013. United States Fish and Wildlife Service (USFWS). 2013. Spartina Invasion and Management, Humboldt Bay. <www.fws.gov/refuge/Humboldt_Bay> Accessed December 14, 2013. Zapiola, M. L., C. K. Campbell, M. D. Butler and C. A. MallorySmith. 2008. Escape and establishment of transgenic glyphosateresistant creeping bentgrass Agrostis stolonifera in Oregon, USA: a 4year study. Journal of Applied Ecology 45.2: 486-494. Zedler, J.B., and S. Kercher. 2005. Wetland resources: Status, trends, ecosystem services, and restorability. Annual Review of Environmental Resources 30: 39-74.

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