Northeastern Naturalist 110 P.P. Daneshgar, et al. 22001177 NORTHEASTERN NATURALIST 2V4(o2l). :2141,0 N–1o2. 02 The Role of Beach Nourishment on the Success of Invasive Asiatic Sand Sedge Pedram P. Daneshgar1,*, Lael B. Phillips1, D. Patrick James1,2, Mitchell G. Mickley1,2, Andrew M. Bohackyj1, 2, Lucas J. Rhoads2, Richard P. Bastian3, and Louise S. Wootton4 Abstract - Beach nourishment, or artificial addition of sediment to a beach, a common practice for rebuilding beaches for recreation and storm protection, can have positive or negative effects on the flora and fauna of dune communities. There has been little work exploring the consequences of nourishment on the impacts of invasive plant species that thrive on dunes. In this study, we explored the impacts of nourishment on the invasiveness of Carex kobomugi (Asiatic Sand Sedge) in secondary dune communities composed primarily of Ammophila breviligulata (American Beachgrass) and Solidago sempervirens (Seaside Goldenrod) at 3 study sites at Island Beach State Park in New Jersey, testing the hypothesis that nourishment promotes sedge invasion. Dune communities were subjected to 5 burial depth treatments in ~15-cm increments ranging from 0 (control) to 60 cm (~24 inches) burial. Growth responses were monitored by quantifying emergent individuals and by harvesting all aboveground biomass at the end of the season. Physiological responses were evaluated using an infrared gas analyzer to quantify photosynthesis rates. Burials lead to a reduction in community diversity and native species biomass, while favoring the invasive species. In addition, Seaside Goldenrod within invaded communities exhibited significantly lower photosynthesis rates than those individuals in non-invaded communities. The results suggest that nourishment will promote Asiatic Sand Sedge invasion to the detriment of native dune species. Introduction A common practice for the maintenance or rebuilding of beaches for recreational and storm protection purposes is nourishment or the artificial addition of sediment to an area that is deficient in sediment. It is well documented that nourishment has localized negative impacts on many coastal species of flora and fauna; burial and habitat reduction can have lethal implications for many populations in the short term (Defeo et al. 2009). Dune plants can be impacted not only by burial itself, but also by soil compaction, which affects gas, nutrient, and water availability in interstitial spaces (Defeo et al. 2009). Another study found that plant biomass decreased by 50% and plant survival decreased by 60% in plots buried under sand (Franks and Peterson 2003). Furthermore, that research found evidence of increased facilitation 1Monmouth University, Marine and Environmental Biology and Policy Program, West Long Branch, NJ 07764. 2Monmouth University, School of Science Summer Research Program, West Long Branch, NJ 07764. 3Monmouth University, Department of Mathematics, West Long Branch, NJ 07764. 4Georgian Court University, Department of Biology, Lakewood, NJ 08701. *Corresponding author - pdaneshg@monmouth.edu. Manuscript Editor: Howard Ginsberg Northeastern Naturalist Vol. 24, No. 2 P.P. Daneshgar, et al. 2017 111 among plants in the burial scenario. Among the buried plots, this facilitation was marked by significantly higher survival rates in plots with higher plant density (Franks and Peterson 2002). Though many studies have reviewed the impacts of nourishment and dredging on macro-invertebrate, shorebird, and fish diversity, none thus far have analyzed the effects of burial on dune plant diversity (Peterson and Bishop 2005). There has been little research on the consequences of nourishment on the impacts of invasive plant species that thrive on dunes. One potential impact of beach nourishment is that the compaction and temporary changes in nutrient, water, and gas availability may lead to the extirpation of invasive species. If this is observed, then nourishment should be encouraged as a management tool to eradicate invasive species followed up by plantings of native species. Conversely, the supplementation of sediment provides new empty niches that may facilitate new invasions (Elton 1958, Levine and D’Antonio 1999, Mack et al. 2000). The disturbance of nourishment alone should be favorable for exotic species invasions, as has been observed in several studies (Lake and Leishman 2004, Sobrino et al. 2002) because disturbances often result in fluctuations of resources, which Davis et al. (2000) proposed is what leads to the invasion of plant communities by exotics. Carex kobomugi Ohwi (Asiatic Sand Sedge), first introduced to New Jersey in 1929, has readily invaded dunes along the east coast from Rhode Island to the Carolinas, largely outcompeting native species and even thriving in dune areas where growth of other plants is limited. The sedge reproduces sexually with separate male and female flowers and asexually through an extensive network of underground rhizomes, which has been observed to increase patches in radius by up to 1 m per year (Wootton 2003, Wootton et al. 2005). Invasion by this species has been shown to decrease mean shoot densities of native species in affected areas (Wootton 2003). Several hypotheses may explain the competitive advantage of Asiatic Sand Sedge. One is that Asiatic Sand Sedge actively removes other species using allelopathic chemicals not otherwise encountered by American dune species (Burkitt and Wootton 2011, Callaway and Ridenour 2004). Another possibility is that North America’s lack of the natural enemies that had limited and suppressed Asiatic Sand Sedge in its native range in Asia, has allowed the species to thrive and spread rapidly here (Maron and Vila 2001). Another explanation comes out of the history of the species’ re-introduction to coastal dunes in the 1970s and 1980s. At the time, Asiatic Sand Sedge was purposely introduced for dune stabilization. The varietal of the species that was used was a hypercompetitive variety that was utilized because it was able to grow successfully (W. Skardek, USDA Cape May Plant Materials Lab, Cape May, NJ, pers. comm.). Although the spread of Asiatic Sand Sedge negatively impacts many native species, it has been successfully used as a dune stabilizer; this history makes removal more difficult and potentially controversial (Wootton 2003). We explored the impacts of nourishment on Asiatic Sand Sedge and associated native dune species. Early observations of researchers at Cape May Plant Materials Laboratory (Shisler et al. 1987), suggest that Asiatic Sand Sedge is relatively Northeastern Naturalist 112 P.P. Daneshgar, et al. 2017 Vol. 24, No. 2 intolerant of burial, while growth of native species, like Ammophila breviligulata Fern. (American Beachgrass) is actually stimulated by active sand accretion. We, however, hypothesize a different effect, which is that burial of dunes invaded by Asiatic Sand Sedge would facilitate its invasion by providing empty niches for the invader to occupy. Field-Site Description Island Beach State Park is a 16-km stretch barrier island located in Ocean County, NJ, bounded by the Atlantic Ocean on the east side and Barnegat Bay on the west side. From east to west on the island, communities transition in the traditional barrier island dune successional pattern from primary dunes to secondary communities to thickets to maritime forest and finally salt marsh. A majority of the Asiatic Sand Sedge invasion in the park occurs in secondary dune communities, which are dominated by American Beachgrass and Solidago sempervirens L. (Seaside Goldenrod) before invasion. We selected 3 study sites that had similar ratios of the 3 study species, American Beachgrass, Seaside Goldenrod, and Asiatic Sand Sedge. The first site (39°53'45.05"N, 74°4'49.86"W), positioned on the front portion of a secondary dune, was within 70–80 m of the mean high water line (MHL) and has been documented as the oldest known colony of Asiatic Sand Sedge in New Jersey (Small 1954). The second site (39°47'25.24"N, 74°5'38.04"W), located at the top of a large secondary dune and within 20 m of a beach access path, lies ~70–90 m from the MHL. The third site (39°47'3.86"N, 74°5'41.47"W), located furthest from the shore at ~100–125 m from the MHL, was bounded by a sand-vehicle access road to the east and thicket communities to the west. Methods In March 2011, we established five 1-m square plots at each site such that there was approximately the same number of the 3 study species (Asiatic Sand Sedge, American Beachgrass, Seaside Goldenrod) within each plot. We conducted a census of all individuals present before the depth of burial was randomly assigned to each plot. Burial depths ranged in 15.2-cm (6-inch) increments from 0 (control) to 60 cm (~24 inches). Each plot was delineated with 4 pieces of rebar ,which we marked in 15-cm increments to allow for accurate addition of sand to the prescribed depth. Each plot was bounded with a sand fence to hold the sand at the burial depth and to reduce movement from wind. We took sand for the burial from pathways adjacent to the dune sites where no vegetation was established. We added sand directly to the plots to the prescribed burial depth, overtopping the individuals growing in them, and then added extra sand around each plot to the prescribed depth to form a 0.5-m buffer around each plot. After the initial nourishment, the prescribed burial depth was maintained through additional additions of sand for the following 3 months due to the fact that the sand initially settled, reducing functional burial depth. After the first 3 months, no more sand was added, and we observed that the depths were stable for the duration of the study . Northeastern Naturalist Vol. 24, No. 2 P.P. Daneshgar, et al. 2017 113 We harvested the aboveground biomass of all plants in each plot in August 2012. The samples were dried for 48 hours in an oven at 65 °C and then weighed. Using those data, we calculated Shannon Wiener diversity indices for each plot. At the peak of the second growing season, in July 2012, we measured lightsaturated photosynthesis rates using an infrared gas analyzer (IRGA) (LI-6400, LiCor Inc., Lincoln, NE) with a photosynthesis photon flux density (PPFD) of 2000 mmol m–2 s–1 and a flow rate of 400 μm/s of CO2 with a reference concentration of 390 ppm. Measurements were taken between the hours of 1100 and 1400 and targeted only fully developed leaves. When an individual leaf did not fully cover the leaf chamber surface (6 cm2), we calculated the area of the individual manually and entered it into the IRGA prior to making a measurement. We also entered the stomatal ratio of each species (1 for Asiatic Sand Sedge and American Beachgrass, 0.5 for Seaside Goldenrod; LiCor Ince. Lincoln, NE) into the IRGA prior to the taking of a reading. We measured light-saturated photosynthesis rates for 3 leaves of 3 different individuals of all 3 of the study species in each plot (9 total measurements per plot). When there were fewer than 3 individuals of a species available, we made measurements of different leaves of the available plants. We also measured the light-saturated photosynthesis rates of 4 American Beachgrass and 4 Seaside Goldenrod individuals picked at random at each site from outside the plots in noninvaded areas adjacent to the study areas for comparison to individuals in invaded areas. These randomly selected individuals were located on the same dune where the plots were located. We calculated descriptive statistics for biomass, diversity, and photosynthesis as mean ± standard deviation. We used a 3 x 5 factorial ANOVA to analyze the randomized complete block design for differences in mean biomass between species, between treatments and their interactions, and a 1-way ANOVA with 5 levels to analyze mean differences in diversity across treatments. There was no effect of block (site) on either of these ANOVA results. We used Tukey’s HSD post-hoc test for mean separation in both ANOVAs. Relationships between photosynthesis, treatment, and species were examined using regression. We declared results as significantly different at α = 0.05 for all tests. All statistical analyses were performed using IBM PASW (formerly SPSS) version 17. Due to the small sample sizes involved, we used non-parametric tests (Kruskall-Wallis and Mann-Whitney) to evaluate differences in overall biomass between the 3 species. Results consistent with the ANOVA and post-hoc tests were obtained when alpha was corrected to 0.05/n. We also used Kruskall-Wallis and Mann-Whitney tests to evaluate the differences in diversity. Results consistent with the 1-way ANOVA were obtained for this test, but the post-hoc tests were no longer significant when using an alpha corrected to 0.05/n. Results The Shannon Wiener diversity of dune communities dropped from 1.034 in the control treatment (no burial) to a value of 0 in the 60-cm burial treatment due to there being only 1 species present in the 60-cm treatment (Asiatic Sand Sedge) at Northeastern Naturalist 114 P.P. Daneshgar, et al. 2017 Vol. 24, No. 2 the time of the harvest (Fig. 1). Every 15 cm of sand added seemed to decrease the diversity of the dune community a little more. The 3 lowest burial treatments (0, 15, and 30 cm burial) did not significantly differ in diversity, but these 3 differed from the 45- and 60-cm burial treatments ( P = 0.009). There was significant difference in aboveground biomass of all 3 study species in all 5 treatments (P < 0.001; Fig. 2). In all treatments, Asiatic Sand Sedge had the greatest aboveground biomass of the 3 species, ranging from 71.6 g/m2 in the 60-cm burial treatment to 98.7 g/m2 in the 15-cm burial treatment. In only the control did Seaside Goldenrod exceed American Beachgrass in aboveground biomass. At 45-cm burial, we observed no Seaside Goldenrod and only 1 g/m2 of American Beachgrass. Neither Seaside Goldenrod nor American Beachgrass were present with 60 cm of burial. The total plot aboveground biomass (all species combined) was marginally different by treatment (P = 0.0792). There was no difference from site to site in the aboveground biomass by treatment ( P = 0.151). Burial did not affect the light-saturated photosynthesis rates of any of the 3 study species (Fig. 3). American Beachgrass light-saturated photosynthesis rates were not affected by the presence or absence of Asiatic Sand Sedge in plots with no burial, while the light-saturated photosynthesis rates of Seaside Goldenrod were apparently reduced by the presence of Asiatic Sand Sedge in plots with no burial (P < 0.001; Fig. 4). Figure 1. Mean Shannon Weiner diversity (+ SD) for the 5 treatment depths across all 3 sites. In the 60-cm treatment, only 1 species was present (Asiatic Sand Sedge) at all 3 sites, resulting in a diversity of zero. Difference letters above the means signify the statistically different means (P = 0.009). Northeastern Naturalist Vol. 24, No. 2 P.P. Daneshgar, et al. 2017 115 Discussion Asiatic Sand Sedge invasion has had detrimental impacts on dunes along the New Jersey coast, spreading at nearly an exponential rate at both Island Beach State Park and Sandy Hook National Recreation Area (Wootton 2003, Wootton et al. 2005). Wootton et al. (2005) also found that dune species diversity declines when Asiatic Sand Sedge is present at Island Beach State Park. Our results indicate that nourishment or burial accelerates species diversity decline associated with invasion, especially when 45 cm of sand or more is added to an invaded dune. This is a relatively low amount of burial compared to the typical nourishments, which often exceed wave-height averages (NRC 1995), or quantities added naturally during storm events. Our finding of Asiatic Sand Sedge’s tolerance to burial is not surprising as it has been shown that the species is very tolerant to sand blasting (Yura and Ogura 2006). In contrast, the tolerance of American Beachgrass to burial is not as clear because studies have yielded mixed results. In California, American Beachgrass was dominant in areas where sand deposition and shoreline change were high (Hacker et al. 2012), and it has been shown that American Beachgrass responds to sand accretion in amounts of up to 50 cm/year with increased shoot vigor and clonal growth (Baye 1990). However, Maun and Lapierre (1984) observed the elimination of populations due to burial. The failure of native species, and in particular American Beachgrass, to respond to burial in this study could be attributed to direct Figure 2. Mean aboveground biomass of the 3 study species by nourishment depth. Asiatic Sand Sedge has significantly more biomass than the other 2 species in all treatments (P less than 0.001), while total plot biomass did not dif fer by treatment (P = 0.0792). Northeastern Naturalist 116 P.P. Daneshgar, et al. 2017 Vol. 24, No. 2 Figure 3. Relationship between burial depth and its impact on mean light-saturated photosynthesis rates of the 3 study species. Missing points are due to absence of the species in the treatment in the second growing season. There was no significance difference in photosynthesis rates by treatment. Figure 4. Mean (+ SD) light-saturated photosynthesis rates for American Beachgrass and Seaside Goldenrod inside and outside the invaded dune areas. Significant differences marked with asterisk (P < 0.001). Northeastern Naturalist Vol. 24, No. 2 P.P. Daneshgar, et al. 2017 117 impacts by the invader. Our photosynthesis results here suggest that Seaside Goldenrod was weakened by the presence of Asiatic Sand Sedge. The success of Asiatic Sand Sedge over native species particularly when buried could be explained by 4 possible mechanisms: better responses to disturbance, ability to outcompete for nutrients, more favorable growth habits, or disruption of vital myccorhizal mutualisms that foster native species growth. The nourishment of dunes provides a disturbance, which may create a more favorable environment for an invader like Asiatic Sand Sedge. In this study, Asiatic Sand Sedge was the only species to emerge from burial depths of over 45 cm after 2 growing seasons. This result may be due to the artificial selection of the Asiatic Sand Sedge in this area, which made it more tolerant to the burial disturbance; thus the response to this disturbance was not surprising. Under conditions of disturbance, some exotic species may become more productive than the native species (Daehler 2003, Hobbs and Humphries 1994) due to a fluctuation of resources (Davis et al. 2000) and new space or niches that were previously unavailable (Elton 1958, Levine and D’Antonio 1999, Mack et al. 2000). The disturbance created here was simply adding sand from an adjacent area where there was no vegetation suggesting that nutrients were low (nutrient content was not measured). Nutrients are generally found in low levels and are limiting to plant growth in dune systems (Maun 1994, Willis 1965). Under limiting conditions, Asiatic Sand Sedge has a greater ability to acquire some nutrients than native plant species. Johnson (2011) observed that Asiatic Sand Sedge was a superior competitor for phosphorus over American Beachgrass, which typically needs the assistance of arbuscular myccorhizal fungi to obtain nutrients. On the Hasaki coastal dunes on the Pacific Ocean, Asiatic Sand Sedge was found to be more effective at obtaining iron and phosphorous under limiting conditions than other native land plants due to the presence of root-endophytic bacteria (Matsuoka et al. 2013). In addition to this, nutrient limitation can also affect a plant’s growth response to burial (Gilbert et al. 2008). The local elimination of American Beachgrass and Seaside Goldenrod due to burial may have been accelerated by even greater nutrient stress caused by Asiatic Sand Sedge. Growth rates of native plant species and Asiatic Sand Sedge also vary greatly. Asiatic Sand Sedge rhizomes are not only deeper than American Beachgrass, but also have a greater number of nodes and internodes (Small 1954), which would suggest a greater ability to establish more root-bearing ramets and occupy more space. Again, this ability allows for the invader to occupy more niches. Rapid growth rates of Asiatic Sand Sedge have been observed and partially explained by the presence of endophytic fungi with the ability to produce gibberellin, a plant hormone, which stimulates growth (Khan et al 2009). These fungi are endophytic, so no previously established fungi are needed for the invader to establish in new sand added to dunes, unlike the natives American Beachgrass and Seaside Goldenrod, which may rely on the presence of arbuscular myccorhizal fungi to successfully establish. Up to 17 different species of arbuscular myccorhizal fungi have been observed to be associated with American Beachgrass-dominated dunes in Massachusetts (Koske and Gemma 1997). It is probable that no spores of arbuscular myccorhizal fungi were present Northeastern Naturalist 118 P.P. Daneshgar, et al. 2017 Vol. 24, No. 2 in the added sand, but this was not tested here; the need for them is essential for the establishment of many dune species (Gemma and Koske 1988). Johnson (2011) reports a negative correlation between arbuscular myccorhizal fungi spore populations and Asiatic Sand Sedge, suggesting that this sedge is reducing the presence of fungal species potentially by the release of fungitoxic root exudates (Li et al. 2009). The exotic-induced reduction in mycchorhizae may lead to nutrient-deficient native species that are less vigorous and thus less able to respond to the disturbance of burial. Further work is needed to definitively explain the me chanisms of Asiatic Sand Sedge invasion success. Our results suggest that burial is not an effective strategy for managing dune invasion, and we recommend that other management strategies should be explored. However, other common management methods also have associated problems. Manual eradication is usually impractical for all but newly established stands due to the deep and extensive root and rhizome systems associated with Asiatic Sand Sedge (Small 1954). Herbicide applications targeting only Asiatic Sand Sedge can be expensive and time consuming (Wootton et al. 2005), and have often not been entirely successful—damage to native species was reduced, but the Asiatic Sand Sedge continued to invade (Wootton et al. 2003). We propose an integrative strategy for managing Asiatic Sand Sedge employing multiple strategies. The process should begin with targeted herbicide applications followed by nourishments and native plantings. American Beachgrass would benefit from the reduction in competition and would fill the new niches provided by the nourishment. Managers should be leery of being too aggressive as non-localized spray applications often have broad effects and can cause high rates of erosion. Dunes populated by Asiatic Sand Sedge are more stable than dunes lacking in vegetation, which are more likely to erode. 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