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Stalter, R. and Lonard, R.I., 2024. Biological flora of sand dunes on the northwest Pacific coastline of North America: Ammophila arenaria (L.) Link and Ammophila breviligulata Fernald. Journal of Coastal Research, 40(5), 994–1000. Charlotte (North Carolina), ISSN 0749-0208. Ammophila arenaria, native to European coastal dunes, was initially introduced on the Pacific coast of North America in 1868, while A. breviligulata was introduced to temperate regions along the Pacific coast in 1935. Both rhizomatous taxa are often dominant on coastal foredunes. Ammophila arenaria has short, thick rhizomes, while A. breviligulata has a spreading rhizome system. Both species have been introduced for sand stabilization and shoreline protection. The primary method of reproduction for both taxa is fragmentation of rhizomes. Ammophila breviligulata aggressively competes with A. arenaria and is now dominant over A. arenaria at many sites. The introduction of A. arenaria and A. breviligulata has contributed to dense cover on foredunes and a concomitant decline of native plants and animals, especially those dependent on open sand habitats. Ammophila breviligulata successfully competes with A. arenaria, which has reduced dune heights, increasing the number of areas vulnerable to storm surges and dune erosion. [ABSTRACT FROM AUTHOR]

Restoration of degraded dunes through revegetation of native dune building species is critical to preserve multiple ecosystem services (i.e., coastal protection, habitat). As dunes exhibit high levels of abiotic stress, it is likely that facilitative interactions among plants occur, reducing stress caused by harsh abiotic conditions, and benefiting restoration efforts. The research was to determine how plant spatial arrangement (clumped vs. dispersed) at the dune slope affects the growth of two dominant dune grasses along the United States Atlantic coast, Ammophila breviligulata (American beachgrass) and Uniola paniculata (sea oats). We determined the local distribution and density of each species through field surveys to inform a manipulative study of planting arrangement and to assess plant growth and survival under different spatial arrangements. Planting arrangement did not influence the survival of U. paniculata, but dispersed plantings had a positive effect on growth parameters (height, stem number, ramet number), compared to the clumped arrangement. No plantings of A. breviligulata survived by the end of the second season, regardless of planting arrangement. Clumped arrangements did not improve growth of U. paniculata or growth and survival of A. breviligulata over the two growing seasons studied; thus, creating clumped arrangements may not be necessary for revegetation efforts. However, observing natural spatial patterns and using this information to guide restoration planting may be beneficial as climate change may impact revegetation results due to increased abiotic stress. Further research focusing on biotic interactions and density of plantings at multiple locations along dunes will contribute to dune restoration planning. [ABSTRACT FROM AUTHOR]

Vegetation shifts can directly alter habitat dynamics and indirectly impact habitat stability relative to disturbance response. Barrier island dune habitats exhibit spatiotemporally dynamic topography that is affected by vegetation. However, vegetation distribution data can be rare and vegetation persistence is largely unknown such that species turnover over in communities can occur unnoticed. This is true despite concerns and documented cases of woody encroachment related to climate change in these ecogeomorphic habitats where physical stability to resist storm erosion varies with vegetation distribution and density. In 1956 and 1957, the vegetation of Island Beach State Park, NJ, was mapped as a permanent record for subsequent ecological study. In June 2020, we remapped the vegetation of 5.4 of 17 km north to south, seaward of the thicket community boundary. We maintained the same classification system as the historic record, physically mapping vegetation patches via GPS. We quantified changes in thicket, heather, and grass community distribution between the two time periods. Habitat persistence and conversion varied in the 63 years. Heath communities saw 80%–97% habitat loss in conversion to woody thicket. Conversely, thicket community distribution drastically increased, replacing heath where it was previously prevalent. This represents the first known documented instances of woody encroachment for Morella pensylvanica. When they did not expand, thicket communities receded landward or underwent turnover to an invasive species. Woody species of interest for dune stabilization occupied areas of similar habitat characteristics. Dune vegetation distribution persistence from 1957 to 2020 was relatively consistent and stable with the exception of heath habitat conversion. Barrier island stability is directly related to vegetation stability such that understanding where community shifts might occur over time can aid in managing and modeling efforts surrounding these dynamic ecogeomorphic habitats. [ABSTRACT FROM AUTHOR]

Abstract Vegetation shifts can directly alter habitat dynamics and indirectly impact habitat stability relative to disturbance response. Barrier island dune habitats exhibit spatiotemporally dynamic topography that is affected by vegetation. However, vegetation distribution data can be rare and vegetation persistence is largely unknown such that species turnover over in communities can occur unnoticed. This is true despite concerns and documented cases of woody encroachment related to climate change in these ecogeomorphic habitats where physical stability to resist storm erosion varies with vegetation distribution and density. In 1956 and 1957, the vegetation of Island Beach State Park, NJ, was mapped as a permanent record for subsequent ecological study. In June 2020, we remapped the vegetation of 5.4 of 17 km north to south, seaward of the thicket community boundary. We maintained the same classification system as the historic record, physically mapping vegetation patches via GPS. We quantified changes in thicket, heather, and grass community distribution between the two time periods. Habitat persistence and conversion varied in the 63 years. Heath communities saw 80%–97% habitat loss in conversion to woody thicket. Conversely, thicket community distribution drastically increased, replacing heath where it was previously prevalent. This represents the first known documented instances of woody encroachment for Morella pensylvanica. When they did not expand, thicket communities receded landward or underwent turnover to an invasive species. Woody species of interest for dune stabilization occupied areas of similar habitat characteristics. Dune vegetation distribution persistence from 1957 to 2020 was relatively consistent and stable with the exception of heath habitat conversion. Barrier island stability is directly related to vegetation stability such that understanding where community shifts might occur over time can aid in managing and modeling efforts surrounding these dynamic ecogeomorphic habitats.

Coastal foredunes form via biophysical feedbacks between sand accretion and burial‐tolerant vegetation and can protect coastlines from hazards such as extreme storms and sea level rise. Predicting how coastal dunes, and the services they provide, will change in the future requires an understanding of the relative roles of the physical and ecological processes that shape their structure and function. Here we assess the relative roles of sand supply, beach morphology, and vegetation in determining foredune morphology, and its change, along a 300‐km stretch of the US Central Atlantic coast. In particular, we used the spatial variability inherent in beaches and dunes of this region to determine the relative importance of shoreline change rate (SCR; a proxy for sand supply to the beach), beach morphology, and grass density of four widespread dune grasses (Uniola paniculata, Ammophila breviligulata, Panicum amarum, and Spartina patens) to foredune morphology metrics (height, width, and aspect ratio) along the North Carolina Outer Banks barrier islands. Foredune morphology and change metrics are correlated with three main factors: multidecadal SCR (1997–2016), beach slope, and dune grass density and species identity. Multidecadal SCR and beach width explained the most variation in, and were positively correlated with, foredune height and width, and were negatively correlated with foredune aspect ratio (height divided by width). In addition, grass density and changes in grass density contributed significantly to foredune morphology change. We found a positive relationship between change in A. breviligulata density and foredune width, which aligns with previous studies on the US Atlantic and Pacific Northwest coasts. Our results demonstrate the interactive roles of beach sand supply and dune grass functional morphology in dune building processes on highly vulnerable coastlines. [ABSTRACT FROM AUTHOR]

Abstract Coastal foredunes form via biophysical feedbacks between sand accretion and burial‐tolerant vegetation and can protect coastlines from hazards such as extreme storms and sea level rise. Predicting how coastal dunes, and the services they provide, will change in the future requires an understanding of the relative roles of the physical and ecological processes that shape their structure and function. Here we assess the relative roles of sand supply, beach morphology, and vegetation in determining foredune morphology, and its change, along a 300‐km stretch of the US Central Atlantic coast. In particular, we used the spatial variability inherent in beaches and dunes of this region to determine the relative importance of shoreline change rate (SCR; a proxy for sand supply to the beach), beach morphology, and grass density of four widespread dune grasses (Uniola paniculata, Ammophila breviligulata, Panicum amarum, and Spartina patens) to foredune morphology metrics (height, width, and aspect ratio) along the North Carolina Outer Banks barrier islands. Foredune morphology and change metrics are correlated with three main factors: multidecadal SCR (1997–2016), beach slope, and dune grass density and species identity. Multidecadal SCR and beach width explained the most variation in, and were positively correlated with, foredune height and width, and were negatively correlated with foredune aspect ratio (height divided by width). In addition, grass density and changes in grass density contributed significantly to foredune morphology change. We found a positive relationship between change in A. breviligulata density and foredune width, which aligns with previous studies on the US Atlantic and Pacific Northwest coasts. Our results demonstrate the interactive roles of beach sand supply and dune grass functional morphology in dune building processes on highly vulnerable coastlines.

Coastal dunes are important protective features against sea level rise and coastal storms. Interactions between dune plant aboveground structures and sediment trapping that allow for dune building and maintenance are well established. More recently, studies documenting belowground biomass for promoting erosion resistance in dominant dune species have been conducted, yet a knowledge gap remains regarding species‐specific characterization of whole plants, specifically with respect to roots, rhizomes, and belowground stems. Our objective was to quantify above‐ and belowground traits of four dominant dune grasses to document the potential for species‐specific effects on dune growth, maintenance, and erosion resistance. We examined above‐ and belowground traits among four prominent dune grasses of the Atlantic and Gulf Coasts of North America: Ammophila breviligulata, Panicum amarum, Spartina patens, and Uniola paniculata. Whole plant samples of each species were collected from the foredune at the US Army Engineer Research and Development Center's Field Research Facility in Duck, North Carolina, USA, and quantified for several above‐ and belowground traits (e.g., stem height, rhizome number and length, root surface area by diameter class, root tensile strength, and mycorrhizal percent infection). Belowground factors known to impact important dune processes, such as rhizome length, mycorrhizal percent infection, and root traits, differed substantially among species. When visualized in multivariate space, all species significantly differed in suites of above‐ and belowground traits. When considering belowground only, Ammophila and Spartina were similar, despite differences in biomass allocation. Species separated along axes related to mycorrhizal association, biomass allocation, and root construction. The four co‐occurring dune grass species were dissimilar in suites of plant traits. Belowground trait differences were driven by those describing root construction, biomass allocation, and mycorrhizal infection. Dissimilarity in above‐ and belowground suites of traits may demonstrate different approaches for surviving the dune environment. Incorporating belowground traits into modeling will enhance predictions of dune response to climate change through interactions between vegetation and dune dynamics that facilitate coastal resistance and resilience. [ABSTRACT FROM AUTHOR]

Abstract Coastal dunes are important protective features against sea level rise and coastal storms. Interactions between dune plant aboveground structures and sediment trapping that allow for dune building and maintenance are well established. More recently, studies documenting belowground biomass for promoting erosion resistance in dominant dune species have been conducted, yet a knowledge gap remains regarding species‐specific characterization of whole plants, specifically with respect to roots, rhizomes, and belowground stems. Our objective was to quantify above‐ and belowground traits of four dominant dune grasses to document the potential for species‐specific effects on dune growth, maintenance, and erosion resistance. We examined above‐ and belowground traits among four prominent dune grasses of the Atlantic and Gulf Coasts of North America: Ammophila breviligulata, Panicum amarum, Spartina patens, and Uniola paniculata. Whole plant samples of each species were collected from the foredune at the US Army Engineer Research and Development Center's Field Research Facility in Duck, North Carolina, USA, and quantified for several above‐ and belowground traits (e.g., stem height, rhizome number and length, root surface area by diameter class, root tensile strength, and mycorrhizal percent infection). Belowground factors known to impact important dune processes, such as rhizome length, mycorrhizal percent infection, and root traits, differed substantially among species. When visualized in multivariate space, all species significantly differed in suites of above‐ and belowground traits. When considering belowground only, Ammophila and Spartina were similar, despite differences in biomass allocation. Species separated along axes related to mycorrhizal association, biomass allocation, and root construction. The four co‐occurring dune grass species were dissimilar in suites of plant traits. Belowground trait differences were driven by those describing root construction, biomass allocation, and mycorrhizal infection. Dissimilarity in above‐ and belowground suites of traits may demonstrate different approaches for surviving the dune environment. Incorporating belowground traits into modeling will enhance predictions of dune response to climate change through interactions between vegetation and dune dynamics that facilitate coastal resistance and resilience.

Forecasting the effects of climate change on the distribution of invasive species can be difficult, because invaders often thrive under novel physical conditions and biotic interactions that differ from those in their native range. In this study, we experimentally examined how rising temperatures and sand burial could alter the abundance and biotic interactions of two invasive beachgrasses, Ammophila arenaria and A. breviligulata, along the U.S. Pacific Northwest coast. We asked whether the current geographic ranges of the two congeners, and thus their effects on dune morphology and coastal ecosystem services, might shift as a consequence of climate driven changes in warming and sand supply. Our results show that A. breviligulata had lower biomass and tiller production when exposed to warming and high rates of sand burial, while A. arenaria showed neutral or positive responses to those treatments. Nevertheless, under all experimental combinations, A. breviligulata had strong negative effects on A. arenaria, while A. arenaria had weaker effects on A. breviligulata. Our models predict that although A. breviligulata mostly excludes A. arenaria, elevated temperatures and high rates of sand burial also increase the likelihood of species coexistence. We suggest that under climate change, the differences in physiological tolerance and the mediation of species interactions could expand the northern distributional limit of A. arenaria but restrict the southern limit of A. breviligulata. Moreover, because beachgrass abundance has direct effects on biophysical functions of dunes, reductions in vigor from warming could alter coastal protection, biodiversity, and carbon sequestration. [ABSTRACT FROM AUTHOR]

Abstract Coastal resilience is threatened as storm‐induced disturbances become more frequent and intense with anticipated changes in regional climate. After severe storms, rapid recovery of vegetation, especially that of dune‐stabilizing plants, is a fundamental property of coastal resilience. Herbivores may affect resilience by foraging and trampling in disturbed areas. Consequently, assessing the impacts of herbivores on recovering vegetation is important for coastal land management. We combined imagery classification, wildlife monitoring, and trend analysis to investigate effects of white‐tailed deer on recovery rates of vegetation four years poststorm in nine overwashed areas. We estimated local deer density with trail cameras, how it relates to an index of primary productivity, and assessed the relationship between deer density and rates of vegetation recovery in overwash fans. Prestorm vegetation cover consisted of shrubs and sporadic patches of beach grass. Poststorm cover was dominated by beach grass. At current rates, vegetation coverage will return to prestorm conditions within the decade, though community transition from grasses to shrubs will take much longer and will vary by site with dune formation. The effect of deer on rates of vegetation recovery was negative, but not statistically significant nor biologically compelling. Although effects of deer trampling on beach grass are evident in classified imagery, deer foraging on beach grass had little effect on its rate of spread throughout overwash fans. While the rate of spread of the primary dune‐building grass was not deleteriously affected by deer, locally high deer densities will likely affect the future establishment and development of herbs and shrubs, which are generally more palatable to deer than beach grass.

The invasive ecosystem engineer Ammophila arenaria, native to Europe, was first introduced to California (USA) in 1896. More than a century later, it has come to dominate coastal foredune vegetation on the west coast of North America to the near exclusion of native species. A. arenaria builds a narrow, steep, peaked, and densely vegetated foredune, in contrast to the broad, more sparsely vegetated foredunes built by the native Elymus mollis. As such, it has modified dune processes by fixing the foredune and disrupting exchange of sediment between the beach, foredune, and dunefield. In the 1930s the congener A. breviligulata, native to the east coast and Great Lakes USA, was first introduced to Oregon, and has been displacing A. arenaria in southern Washington. Ammophila spp. have drastically reduced biodiversity, outcompeting native plant species, and displacing both invertebrate and vertebrate species. Restoration of west coast dunes through the removal of Ammophila began in the 1990s. Methods usually consist of one or a combination of manual digging, burning/herbicides, or excavation with heavy equipment. There are benefits and disadvantages to each method. Manual removal has proven most effective at restoring foredune form and process but is expensive. Excavation and herbicides may result in the loss of foredune morphology. Managers must articulate goals carefully before selecting restoration methods.

The microbial community composition of coastal dunes can vary across environmental gradients, with the potential to impact erosion and deposition processes. In coastal foredunes, invasive plant species establishment can create and alter environmental gradients, thereby altering microbial communities and other ecogeomorphic processes with implications for storm response and management and conservation efforts. However, the mechanisms of these processes are poorly understood. To understand how changing microbial communities can alter these ecogeomorphic dynamics, one must first understand how soil microbial communities vary as a result of invasion. Towards this goal, bacterial communities were assessed spatially along foredune microhabitats, specifically in barren foredune toe and blowout microhabitats and in surrounding vegetated monocultures of native Ammophila breviligulata and invasive Carex kobomugi. Across dune microhabitats, microbial composition was more dissimilar in barren dune toe and blowout microhabitats than among the two plant species, but it did not appear that it would favor the establishment of one plant species over the other. However, the subtle differences between the microbial community composition of two species could ultimately aid in the success of the invasive species by reducing the proportions of bacterial genera associated exclusively with A. breviligulata. These results suggest that arrival time may be crucial in fostering microbiomes that would further the continued establishment and spread of either plant species.

Coastal resilience is threatened as storm‐induced disturbances become more frequent and intense with anticipated changes in regional climate. After severe storms, rapid recovery of vegetation, especially that of dune‐stabilizing plants, is a fundamental property of coastal resilience. Herbivores may affect resilience by foraging and trampling in disturbed areas. Consequently, assessing the impacts of herbivores on recovering vegetation is important for coastal land management.We combined imagery classification, wildlife monitoring, and trend analysis to investigate effects of white‐tailed deer on recovery rates of vegetation four years poststorm in nine overwashed areas. We estimated local deer density with trail cameras, how it relates to an index of primary productivity, and assessed the relationship between deer density and rates of vegetation recovery in overwash fans.Prestorm vegetation cover consisted of shrubs and sporadic patches of beach grass. Poststorm cover was dominated by beach grass. At current rates, vegetation coverage will return to prestorm conditions within the decade, though community transition from grasses to shrubs will take much longer and will vary by site with dune formation.The effect of deer on rates of vegetation recovery was negative, but not statistically significant nor biologically compelling. Although effects of deer trampling on beach grass are evident in classified imagery, deer foraging on beach grass had little effect on its rate of spread throughout overwash fans.While the rate of spread of the primary dune‐building grass was not deleteriously affected by deer, locally high deer densities will likely affect the future establishment and development of herbs and shrubs, which are generally more palatable to deer than beach grass. [ABSTRACT FROM AUTHOR]

Coastal foredunes provide the first line of defense against rising sea levels and storm surge and for this reason there is increasing interest in understanding and modeling foredune formation and post‐storm recovery. However, there is limited observational data available to provide empirical guidance for the development of model parameterizations. To provide guidance for improved representation of dune grass growth in models, we conducted a two‐year multi‐species transplant experiment on Hog Island, VA, U.S.A. and measured the dependence of plant growth on elevation and distance from the shoreline, as well as the relationship between plant growth and sand accumulation. We tracked total leaf growth (length) and aboveground leaf length and found that Ammophila breviligulata (American beachgrass) and Uniola paniculata (sea oats) grew more than Spartina patens (saltmeadow cordgrass) by a factor of 15% (though not statistically significant) and 45%, respectively. Our results also suggest a range of basal/frontal area ratios (an important model parameter) from 0.5‐1 and a strong correlation between transplant growth and total sand deposition for all species at the scale of two years, but not over shorter temporal scales. Distance from the shoreline and elevation had no effect on transplant growth rate but did have an effect on survival. Based on transplant survival, the seaward limit of vegetation at the end of the experiment was approximately 30 m from the MHWL and at an elevation of 1.43 m, corresponding to inundation less than 7.5% of the time according to total water level calculations. Results from this experiment provide evidence for the dune‐building capacity of all three species, suggesting S. patens is not a maintainer species, as previously thought, but rather a moderate dune builder even though its growth is less stimulated by sand deposition than A. breviligulata and U. paniculata. © 2019 John Wiley & Sons, Ltd. [ABSTRACT FROM AUTHOR]

Aims: Soil communities are shaped by multiple biotic and environmental components. We assessed the role of multiple factors in shaping soil bacterial and nematode communities in an ongoing plant invasion in western Michigan sand dunes. We aimed to determine which factors were important for structuring soil communities.Methods: We surveyed seven sites containing the exotic plant, Leymus arenarius, in western Michigan, USA sand dunes. We sampled the plant community and collected soil cores to assess bacterial and nematode communities. We assessed bacterial functional diversity using Biolog Ecoplates. Nematode communities were assessed based on morphological identification.Results: Temperature was a strong determinant of bacterial functional diversity and community composition. Precipitation and Ammophila density were also correlated with bacterial function composition. Nematode communities were shaped by a combination of temperature, precipitation, Ammophila density, and to a lesser degree soil organic matter and Leymus density. Soil bacteria and bacterivorous nematode assemblages were not significantly associated in their responses to the environment.Conclusions: Soil communities of bacteria and nematodes in this early successional dune system are affected strongly by changes in climatic factors, rather than exotic plant presence. [ABSTRACT FROM AUTHOR]

Previous work on the US Atlantic coast has generally shown that coastal foredunes are dominated by two dune grass species, Ammophila breviligulata (American beachgrass) and Uniola paniculata (sea oats). From Virginia northward, A. breviligulata dominates, while U. paniculata is the dominant grass south of Virginia. Previous work suggests that these grasses influence the shape of coastal foredunes in species-specific ways, and that they respond differently to environmental stressors; thus, it is important to know which species dominates a given dune system. The range boundaries of these two species remains unclear given the lack of comprehensive surveys. In an attempt to determine these boundaries, we conducted a literature survey of 98 studies that either stated the range limits and/or included field-based studies/observations of the two grass species. We then produced an interactive map that summarizes the locations of the surveyed papers and books. The literature review suggests that the current southern range limit for A. breviligulata is Cape Fear, NC, and the northern range limit for U. paniculata is Assateague Island, on the Maryland and Virginia border. Our data suggest a northward expansion of U. paniculata, possibly associated with warming trends observed near the northern range limit in Painter, VA. In contrast, the data regarding a range shift for A. breviligulata remain inconclusive. We also compare our literature-based map with geolocated records from the Global Biodiversity Information Facility and iNaturalist research grade crowd-sourced observations. We intend for our literature-based map to aid coastal researchers who are interested in the dynamics of these two species and the potential for their ranges to shift as a result of climate change.

ABSTRACT Day, F.P.; Adams, E.C.; Gibala-Smith, L.A.; Graziani, D.J.; McMillan, B.; Sedghi, N.; Shafer, J., and Smith, M., 2018. Determining change in coastal barrier island dune vegetation following a decade of nitrogen fertilization. Nitrogen deposition from agricultural and industrial sources is a threat to terrestrial biodiversity, and impacts are likely to be greatest in nitrogen-deficient systems. The results of a 10 year fertilization treatment on nitrogen-deficient coastal barrier island dunes and the decade following cessation of fertilizer application are reported here. The study objective was to determine the resilience of the dune plant communities. Would these communities return to their former state in terms of density, species composition, and species diversity after nitrogen additions ended, and if so, how rapid was their recovery? Plant density by species and species diversity were determined over a 20 year period from permanent plots established on three different-aged dunes on a Virginia barrier island. Two dominant species responded differently to nitrogen additions. Ammophila breviligulata abundance was enhanced by the additions, and, in most cases, Spartina patens abundance was repressed. On the older Hog Island dunes, the addition of nitrogen over a 10 year period had a dramatic and rapid negative effect on diversity. However, recovery to diversity values observed in the control plots was rapid, beginning almost immediately after nitrogen additions were stopped. The dune vegetation on a Virginia barrier island appears to be quite resilient, as it rapidly recovered from diversity loss after 10 years of nitrogen additions. [ABSTRACT FROM AUTHOR]

Previous work on the US Atlantic coast has generally shown that coastal foredunes are dominated by two dune grass species, Ammophila breviligulata (American beachgrass) and Uniola paniculata (sea oats). From Virginia northward, A. breviligulata dominates, while U. paniculata is the dominant grass south of Virginia. Previous work suggests that these grasses influence the shape of coastal foredunes in species-specific ways, and that they respond differently to environmental stressors; thus, it is important to know which species dominates a given dune system. The range boundaries of these two species remains unclear given the lack of comprehensive surveys. In an attempt to determine these boundaries, we conducted a literature survey of 98 studies that either stated the range limits and/or included field-based studies/observations of the two grass species.We then produced an interactive map that summarizes the locations of the surveyed papers and books. The literature review suggests that the current southern range limit for A. breviligulata is Cape Fear, NC, and the northern range limit for U. paniculata is Assateague Island, on the Maryland and Virginia border. Our data suggest a northward expansion of U. paniculata, possibly associated with warming trends observed near the northern range limit in Painter, VA. In contrast, the data regarding a range shift for A. breviligulata remain inconclusive. We also compare our literature-based map with geolocated records from the Global Biodiversity Information Facility and iNaturalist research grade crowd-sourced observations. We intend for our literature-based map to aid coastal researchers who are interested in the dynamics of these two species and the potential for their ranges to shift as a result of climate change. [ABSTRACT FROM AUTHOR]

Sand dunes are important geomorphic formations of coastal ecosystems that are critical in protecting human populations that live in coastal areas. Dune formation is driven by ecomorphodynamic interactions between vegetation and sediment deposition. While there has been extensive research on responses of dune grasses to sand burial, there is a knowledge gap in understanding mechanisms of acclimation between similar, coexistent, dune-building grasses such as Ammophila breviligulata (C3), Spartina patens (C4), and Uniola paniculata (C4). Our goal was to determine how physiological mechanisms of acclimation to sand burial vary between species. We hypothesize that (1) in the presence of burial, resource allocation will be predicated on photosynthetic pathway and that we will be able to characterize the C3 species as a root allocator and the C4 species as leaf allocators. We also hypothesize that (2) despite similarities between these species in habitat, growth form, and life history, leaf, root, and whole plant traits will vary between species when burial is not present. Furthermore, when burial is present, the existing variability in physiological strategy will drive species-specific mechanisms of survival. In a greenhouse experiment, we exposed three dune grass species to different burial treatments: 0 cm (control) and a one-time 25-cm burial to mimic sediment deposition during a storm. At the conclusion of our study, we collected a suite of physiological and morphological functional traits. Results showed that Ammophila decreased allocation to aboveground biomass to maintain root biomass, preserving photosynthesis by allocating nitrogen (N) into light-exposed leaves. Conversely, Uniola and Spartina decreased allocation to belowground production to increase elongation and maintain aboveground biomass. Interestingly, we found that species were functionally distinct when burial was absent; however, all species became more similar when treated with burial. In the presence of burial, species utilized functional traits of rapid growth strategy, although mechanisms of change were interspecifically variable. [ABSTRACT FROM AUTHOR]

Vegetation and biogeomorphology are highly coupled in beach dune systems, but plant species effects on abating storm erosion are largely unexplored., We quantified coastal dune erosion from Hurricane Sandy (October 2012) as a function of pre-storm system characteristics - dune height, beach width and dominant vegetation stabilizing dunes (native Ammophila breviligulata or invasive Carex kobomugi) at Island Beach State Park, New Jersey, USA. We assessed dune erosion using a combination of pre- and post-Sandy aerial image spatial analyses in ArcGIS and GPS field mapping. Our two erosion metrics are novel, macroscale 2D surface area changes and Dune Crest Transgression, the later of which is measured at the microscale (1 m−1) and analysed using a mixed model incorporating spatial autocorrelation., We document a species-specific effect on collision erosion. Although C. kobomugi reduces native diversity and abundance, it may be beneficial for coastal protection, as dunes fronted and stabilized by C. kobomugi suffered less erosion than those dominated by A. breviligulata under the same abiotic conditions., Dune height and beach width were equal for species prior to the storm and therefore do not account for or confound differences in erosion. Similarly, traditional calculations of erosion with volumetric loss confer these results., Synthesis and applications. This is the first study to show a species effect on coastal dune erosion. Native Ammophila breviligulata stabilized dune stretches suffered more erosion from Hurricane Sandy than complimentary invasive Carex kobomugi stretches, contradicting anecdotal reports that foredunes stabilized by a shorter statured species are more prone to erosion. This study highlights the importance of vegetation for dune stability and management using two novel metrics for erosion. Our erosion metrics are related to volumetric loss, can be monitored and calculated by managers with or without remote sensing, and can be applied to other systems. Discussions on coastal management of dunes as habitats and protective buffers must include vegetation and the results of this study suggest that not all species are equal with regard to their ability to combat storm erosion. Multidisciplinary studies with applied implications will grow increasingly important as storms continue to grow more frequent, severe and unpredictable with climate change. [ABSTRACT FROM AUTHOR]

Understanding interactions between above- and belowground components of ecosystems is an important next step in community ecology. These interactions may be fundamental to predicting ecological responses to global change because indirect effects occurring through altered species interactions can outweigh or interact with the direct effects of environmental drivers. In a multiyear field experiment (2010-2015), we tested how experimental addition of a mutualistic leaf endophyte (Epichloë amarillans) associated with American beachgrass (Ammophila breviligulata) interacted with an altered precipitation regime (±30%) to affect the belowground microbial community. Epichloë addition increased host root biomass at the plot scale, but reduced the length of extraradical arbuscular mycorrhizal (AM) fungal hyphae in the soil. Under ambient precipitation alone, the addition of Epichloë increased root biomass per aboveground tiller and reduced the diversity of AM fungi in A. breviligulata roots. Furthermore, with Epichloë added, the diversity of root-associated bacteria declined with higher soil moisture, whereas in its absence, bacterial diversity increased with higher soil moisture. Thus, the aboveground fungal mutualist not only altered the abundance and composition of belowground microbial communities but also affected how belowground communities responded to climate, suggesting that aboveground microbes have potential for cascading influences on community dynamics and ecosystem processes that occur belowground. [ABSTRACT FROM AUTHOR]

Barrier islands are at the forefront of storms and sea-level rise. High disturbance regimes and sediment mobility make these systems sensitive and dynamic. Island foredunes are protective structures against storm-induced overwash that are integrally tied to dune grasses via biogeomorphic feedbacks. Shifts in dune grass dominance could influence dune morphology and susceptibility to overwash, altering island stability. In a glasshouse study, two dune grasses, Ammophila breviligulata and Uniola paniculata, were planted together and subjected to a 20 cm burial to quantify morphological and physiological responses. Burial had positive effects on both plants as indicated by increased electron transport rate and total biomass. Ammophila breviligulata performance declined when planted with U. paniculata. Uniola paniculata was not affected when planted with A. breviligulata but did have higher water use efficiency and nitrogen use efficiency. Planted in mixture, differential reallocation of biomass occurred between species potentially altering resource acquisition further. As U. paniculata migrates into A. breviligulata dominated habitat and A. breviligulata performance diminishes, biotic interactions between these and other species may affect dune formation and community structure. Our study emphasizes the importance of studying biotic interactions alongside naturally occurring abiotic drivers. [ABSTRACT FROM AUTHOR]

Coastal regions are inherently and increasingly vulnerable and geomorphologically unstable, yet are invaluable economic and residential hubs. Dunes are dynamic buffers to erosion and the most natural, economical, and effective defense for coastal communities. Vegetation is integral to dune structure as it facilitates accretion and stabilization. Differences in the vegetation and root density likely translate to variability in coastal erosion prevention, but this notion has been largely unconsidered. We directly compared stabilizing factors, depth and density, of the root systems of two dominant mid-Atlantic dune plant species, native American beach grass ( Ammophila breviligulata) and invasive Asiatic sand sedge ( Carex kobomugi ). Despite high plant density, C. kobomugi is targeted for removal in restoration efforts as its roots are assumed to provide less effective stabilization than A. breviligulata . We collected 30 cores and hand dug 14 A. breviligulata ramets at Island Beach State Park, New Jersey to examine biomass, root:shoot ratios, and root density. C. kobomugi had a more extensive root system with a root:shoot ratio of 11.36:1 compared to 1.62:1 for A. breviligulata . Similarly, cores 60 cm deep and 7.6 cm wide were sufficient to attain fully intact A. breviligulata roots, which did not extend deeper than 40 cm, but insufficient for C. kobomugi roots which extended beyond the sampling system vertically and horizontally. Scaling these findings to m −2 , aboveground biomass is relatively equal, but C. kobomugi had over 700% more root mass m −2 than A. breviligulata. These results have strong implications for dune management. The root system of C. kobomugi may be better adapted to stabilize dunes and thus protect coastal areas during small and large-scale perturbations than previously supposed. This is a unique situation whereby the creation of monocultures will hyperstabilize dunes and make them more resistant to erosion at the cost of reduced biodiversity within the framework of resiliency. [ABSTRACT FROM AUTHOR]

For many ecosystems, one of the primary avenues of climate impact may be through changes to foundation species, which create habitats and sustain ecosystem services. For plants, microbial symbionts can often act as mutualists under abiotic stress and may mediate foundational plant responses to climate change. We manipulated the presence of endophytes in Ammophila breviligulata, a foundational sand dune species, to evaluate their potential to influence plant responses to climate change. We simulated projected climate change scenarios for temperature and precipitation using a growth chamber experiment. A 5 °C increase in temperature relative to current climate in northern Michigan reduced A. breviligulata survival by 45 %. Root biomass of A. breviligulata, which is critical to dune stabilization, was also strongly reduced by temperature. Plants inoculated with the endophyte had 14 % higher survival than endophyte-free plants. Contrary to our prediction, endophyte symbiosis did not alter the magnitude or direction of the effects of climate manipulations on A. breviligulata survival. However, in the absence of the endophyte, an increase in temperature increased the number of sand grains bound by roots by 80 %, while in symbiotic plants sand adherence did not significantly respond to temperature. Thus, plant–endophyte symbiosis actually negated the benefits in ecosystem function gained under a warmer climate. This study suggests that heat stress related to climate change in the Great Lakes may compromise the ability of A. breviligulata to stabilize dune ecosystems and reduce carbon storage and organic matter build-up in these early-successional systems due to reduced plant survival and root growth. [ABSTRACT FROM AUTHOR]

The spatial distribution of species is mediated by a combination of biotic interactions and environmental conditions. Understanding the relative importance of these factors and how they interact is particularly important for predicting the spread of non-native species and their impact on resident communities., We used a 3-species Lotka- Volterra model parameterized with field and experimental data to understand the potential for continued spread by an introduced, non-native, dune-building beach grass ( Ammophila breviligulata) and whether this invasion will result in coexistence or displacement in the resident beach grass communities (native Elymus mollis and introduced, non-native Ammophila arenaria) of the US Pacific Northwest., We also used the model to investigate to what extent different rates of ocean-driven sand supply mediate the outcomes of beach grass species interactions., Our model suggests that A. breviligulata could invade and dominate dunes across the range of sand supply rates observed in the region. We found that sand supply influenced intra- vs. interspecific interactions, the strength of positive and indirect effects among beach grasses and the long-term abundances of the beach grass species themselves., Of the two non-natives, A. breviligulata is the inferior dune-building species. Thus, our results suggest that further invasions by A. breviligulata could reduce the coastal protection services afforded by tall dunes currently dominated by A. arenaria., Synthesis: In systems with strong environmental forcing and stressful conditions such as coastal dunes, environmentally mediated positive and indirect species interactions can govern invasion success and long-term native-non-native coexistence. In doing so, these interactions ultimately shape community structure and ecosystem function. Understanding the joint effects of environmental forcing and species interactions on community assembly is particularly important in cases where species introductions can alter ecosystem services, such as coastal protection, which are vulnerable to the effects of climate change. [ABSTRACT FROM AUTHOR]

Plant biodiversity is being lost at a rapid rate. This has spurred much interest in elucidating the consequences of this loss for higher trophic levels. Experimental tests have shown that both plant species diversity and genetic diversity within a plant species can influence arthropod community structure. However, the majority of these studies have been conducted in separate systems, so their relative importance is currently unresolved. Furthermore, potential interactions between the two levels of diversity, which likely occur in natural systems, have not been investigated. To clarify these issues, we conducted three experiments in a freshwater sand dune ecosystem. We (1) independently manipulated plant species diversity, (2) independently manipulated genetic diversity within the dominant plant species, Ammophila breviligulata, and (3) jointly manipulated genetic diversity within the dominant plant and species diversity. We found that genetic diversity within the dominant plant species, Ammophila breviligulata, more strongly influenced arthropod communities than plant species diversity, but this effect was dependent on the presence of other species. In species mixtures, A. breviligulata genetic diversity altered overall arthropod community composition, and arthropod richness and abundance peaked at the highest level of genetic diversity. Positive nonadditive effects of diversity were detected, suggesting that arthropods respond to emergent properties of diverse plant communities. However, in the independent manipulations where A. breviligulata was alone, effects of genetic diversity were weaker, with only arthropod richness responding. In contrast, plant species diversity only influenced arthropods when A. breviligulata was absent, and then only influenced herbivore abundance. In addition to showing that genetic diversity within a dominant plant species can have large effects on arthropod community composition, these results suggest that understanding how species diversity and genetic diversity interact to influence community structure may be critically important for predicting the consequences of biodiversity loss. [ABSTRACT FROM AUTHOR]

Both plant species diversity and genetic diversity within a plant species can affect community properties and ecosystem processes. However, the relative contribution of species diversity and genetic diversity to ecosystem functioning is poorly known. Furthermore, ecosystem processes may respond non-additively to interactions between species diversity and genetic diversity. If interactive effects exist, the impact of biodiversity loss may not be predictable from simple assessments of either species diversity or genetic diversity alone., Here, we addressed how plant species diversity and genetic diversity within a dominant species independently and interactively influenced plant community biomass in a Great Lakes sand dune ecosystem. To test the independent effects of diversity, we established two experiments. In one, we manipulated genetic diversity within the dominant dune species, Ammophila breviligulata. In the other, we manipulated the number of plant species, excluding A. breviligulata. Then, to test for interactive effects, we constructed communities that varied the number of species and levels of genetic diversity within A. breviligulata., Although there were no independent effects of either species diversity or genetic diversity within A. breviligulata on biomass production in this system, interactive effects of species diversity and genetic diversity significantly influenced overall above-ground biomass production of the plant communities. Specifically, as genetic diversity within the dominant species increased, the relationship between species diversity and community-level biomass shifted from negative to positive. Negative non-additive effects of diversity drove this pattern., Synthesis. These results show, for the first time, that interactions between plant species diversity and genetic diversity within a dominant species can alter biomass production, highlighting the importance of incorporating interactions between levels of biodiversity into our understanding of how biodiversity influences ecosystem function. [ABSTRACT FROM AUTHOR]

Vegetation at the aquatic-terrestrial interface can alter landscape features through its growth and interactions with sediment and fluids. Even similar species may impart different effects due to variation in their interactions and feedbacks with the environment. Consequently, replacement of one engineering species by another can cause significant change in the physical environment. Here we investigate the species-specific ecological mechanisms influencing the geomorphology of U.S. Pacific Northwest coastal dunes. Over the last century, this system changed from open, shifting sand dunes with sparse vegetation (including native beach grass, Elymus mollis), to densely vegetated continuous foredune ridges resulting from the introduction and subsequent invasions of two nonnative grass species (Ammophila arenaria and Ammophila breviligulata), each of which is associated with different dune shapes and sediment supply rates along the coast. Here we propose a biophysical feedback responsible for differences in dune shape, and we investigate two, non-mutually exclusive ecological mechanisms for these differences: (1) species differ in their ability to capture sand and (2) species differ in their growth habit in response to sand deposition. To investigate sand capture, we used a moveable bed wind tunnel experiment and found that increasing tiller density increased sand capture efficiency and that, under different experimental densities, the native grass had higher sand capture efficiency compared to the Ammophila congeners. However, the greater densities of nonnative grasses under field conditions suggest that they have greater potential to capture more sand overall. We used a mesocosm experiment to look at plant growth responses to sand deposition and found that, in response to increasing sand supply rates, A. arenaria produced higher-density vertical tillers (characteristic of higher sand capture efficiency), while A. breviligulata and E. mollis responded with lower-density lateral tiller growth (characteristic of lower sand capture efficiency). Combined, these experiments provide evidence for a species-specific effect on coastal dune shape. Understanding how dominant ecosystem engineers, especially nonnative ones, differ in their interactions with abiotic factors is necessary to better parameterize coastal vulnerability models and inform management practices related to both coastal protection ecosystem services and ecosystem restoration. [ABSTRACT FROM AUTHOR]

Beach erosion problems have become prevalent in Korea as a result of its economic growth and coastal development. This paper summarizes a comprehensive approach used to mitigate beach erosion that occurred after the construction of a coastal road with a seawall along Namae Beach in Korea. Field measurements of waves and currents were performed to identify the wave-induced current pattern associated with the observed shoreline erosion in front of the seawall. The construction of a short jetty and two submerged reef mounds was proposed to reduce the wave-induced current and resulting beach erosion. After the jetty construction from May to August, 2008, the shoreline advanced offshore continuously in the zone affected by the jetty. Field monitoring will be continued during the reef mound construction and for beach nourishment proposed to maintain a wide beach in front of the seawall. This mitigation project indicates that the effect of a coastal road on an adjacent beach should be taken into account before its siting because of the long-term evolution of the beach morphology. [ABSTRACT FROM AUTHOR]

To provide necessary information for the management of biodiversity on Sable Island, Nova Scotia, we studied the effects of feral horses on vegetation using exciosures and ancillary observations. Nine plant communities inside and outside of six exclosures were compared using various vegetation parameters and Mann-Whitney tests to evaluate the significance of dif- ferences. The most important findings were as follows: (1) effects of horses were greatest in the Marram (Ammophila bre- viligulata) grassland and much less in the communities that were not dominated by Marram Grass; (2) effects on Marram grassland varied substantially among sites; (3) the cover of standing litter of herbaceous plants was on average of 9.3 times greater inside exclosures in grassland habitats; (4) the cover of living foliage was usually higher inside exclosures, but not all differences were significant; (5) species richness and species diversity were not substantially affected; (6) the average cover of Marram Grass, the most abundant plant and a key sand binder on the island, was greater inside exclosures in six of seven study sites, significantly so in three of them; and (7) there were inconsistent differences in cover of other species at different sites. Wetland habitats cover a relatively small portion of Sable Island, but they support much of the plant biodiver- sity. There is evidence of strong but variable effects of horses on wetland vegetation. "Horse lawns" are littoral habitats dominated by Agrostis stolonifera (Carpet Bentgrass) and other low-growing plants. The lawn habitats represent less than 1% of the island's vegetation, and their presence is believed to be due to grazing and trampling by horses. [ABSTRACT FROM AUTHOR]

Because of the economic and environmental importance of stabilizing fragile sand dune habitats, restoration of dunes has become a common practice. Restoration efforts in the Great Lakes and East Coast regions of North America often consist of planting monocultures of the dominant native grass species, Ammophila breviligulata. We evaluated 18 dune restoration projects in the Great Lakes region conducted over the past 25 years. We characterized attributes of diversity (plants and insects), vegetation structure (plant biomass and cover), and ecological processes (soil nutrients and mycorrhizal fungi abundance) in each restoration, and we compared these measures to geographically paired natural dune communities. Restoration sites were similar to reference sites in most measured variables. Differences between restorations and reference sites were mostly explained by differences in ages, with the younger sites supporting slightly lower plant diversity and mycorrhizal spore abundance than older sites. Plant community composition varied little between restored and reference sites, with only one native forb species, Artemisia campestris, occurring significantly more often in reference sites than restored sites. Although it remains unclear whether more diverse restoration plantings could accelerate convergence on the ecological conditions of reference dunes, in general, traditional restoration efforts involving monoculture plantings of A. breviligulata in Great Lakes sand dunes appear to achieve ecological conditions found in reference dunes. [ABSTRACT FROM AUTHOR]

The distribution and diversity of Pratylenchus species associated with Ammophila arenaria was investigated in its natural range of distribution. Twelve localities with vigorous stands of A. arenaria along the European Atlantic and Mediterranean coasts were sampled. The populations were identified based on morphology and morphometrics, and further characterised based on sequences of the rDNA D2D3 region. Pratylenchus spp. were present in all of the sampled sites. A total of 19 populations were detected belonging to Pratylenchus dunensis, P. brzeskii, P. pratensis or P. penetrans. Pratylenchus dunensis was widely distributed from Blakeney Point (UK) to Comporta (Portugal). Pratylenchus brzeskii was found in South European localities along the Atlantic coast and also in the Mediterranean region. Pratylenchus pratensis was found associated with A. arenaria for the first time and occurred at different locations along the Atlantic coast. Pratylenchus penetrans was only detected in Biarritz (France). The P. dunensis populations from the south west Iberian Peninsula differed from the original P. dunensis description and showed two incisures on the lip region instead of one. Pratylenchus brzeskii populations did not vary morphologically from the original descriptions; however, the range of their morphometrical characters was wider than that of the type population. The D2D3 rDNA region revealed large interspecific and low intraspecific variation, supporting the morphological identification. The phylogenetic relationships of the populations with respect to other species of the genus were inferred from partial sequences of the rDNA and positioned P. dunensis within the same group as P. convallariae, P. penetrans and P. fallax. [ABSTRACT FROM AUTHOR]

Topography and vegetation of restored dunes on a developed barrier island were examined after a large-scale beach nourishment project. Restoration began in 1993 using sand-trapping fences and Ammophila breviligulata Fern. plantings. Subsequent growth of dunes was favored by installing new fences and suspending beach raking to accommodate nesting birds. Plant species richness, percent cover of vegetation, and height of A. breviligulata were sampled in 1999 on seven shore perpendicular transects in six dune microhabitats (backdune, primary crest, mid-foredune, swale, seaward-most fenced ridge, incipient dune on the backbeach). A total of 26 plant taxa were found at all seven sites. Richness and percent cover were greatest in the backdune and crest, especially in locations that predated the 1992 nourishment. Richness was greater where fences enhanced stabilization. Fences initially compensate for time and space and allow vegetation to develop rapidly, but maintenance nourishment is required to protect against wave erosion and ensure long-term viability of habitat. An expanded environmental gradient is an option, where beach nourishment provides space for a species-rich crest and backdune to develop, while the incipient dune remains dynamic. Options where space is restricted include a dynamic, full-sized seaward section of a naturally functioning dune (truncated gradient) or a spatially restricted sampler of a wider natural dune (compressed gradient) maintained using fences. Expanded and truncated gradients may become self-sustaining and provide examples of natural cycles of change. Compressed gradients provide greater species richness and flood protection for the available space, but habitats are vulnerable to erosion, and resident views may be impaired. [ABSTRACT FROM AUTHOR]

Only recently has the importance of positive interactions among plant species in structuring natural communities been supported by experimental evidence. Most studies have focused on interactions between a pair of species at a single life-history stage. In this study positive interactions between a woody nitrogen-fixing shrub (Myrica pensylvanica) and two herbaceous sand dune species (Solidago sempervirens, Ammophila breviligulata) which frequently grow beneath shrub canopies are examined throughout the life cycles of the herbaceous species. Comparisons of S. sempervirens and A. breviligulata growing beneath and outside M. pensylvanica shrubs show that plants growing in association with shrubs are larger, are more likely to flower, produce greater numbers of flowers and seeds, have higher midday xylem water potentials, have higher tissue nitrogen concentrations, and have higher photosynthetic efficiencies. Measurements of environmental conditions show that areas beneath shrubs are more shaded, have lower soil temperatures, and have higher soil nitrogen levels. The results from experimental manipulations designed to test the effects of Myrica shrubs on understory species suggest that the observed differences in plant performance are strongly influenced by canopy shading and soil nutrient enrichment associated with the shrubs. The results demonstrate that M. pensylvanica facilitates growth, reproduction, and recruitment of S. sempervirens and A. breviligulata growing beneath it. This study, one of the few to examine positive interactions at different life-history stages, supports previous predictions that positive interactions may be particularly important in plant communities characterized by physiologically stressful conditions. [ABSTRACT FROM AUTHOR]

The invasive ecosystem engineer Ammophila arenaria, native to Europe, was first introduced to California (USA) in 1896. More than a century later, it has come to dominate coastal foredune vegetation on the west coast of North America to the near exclusion of native species. A. arenaria builds a narrow, steep, peaked, and densely vegetated foredune, in contrast to the broad, more sparsely vegetated foredunes built by the native Elymus mollis. As such, it has modified dune processes by fixing the foredune and disrupting exchange of sediment between the beach, foredune, and dunefield. In the 1930s the congener A. breviligulata, native to the east coast and Great Lakes USA, was first introduced to Oregon, and has been displacing A. arenaria in southern Washington. Ammophila spp. have drastically reduced biodiversity, outcompeting native plant species, and displacing both invertebrate and vertebrate species. Restoration of west coast dunes through the removal of Ammophila began in the 1990s. Methods usually consist of one or a combination of manual digging, burning/herbicides, or excavation with heavy equipment. There are benefits and disadvantages to each method. Manual removal has proven most effective at restoring foredune form and process but is expensive. Excavation and herbicides may result in the loss of foredune morphology. Managers must articulate goals carefully before selecting restoration methods. [ABSTRACT FROM AUTHOR]