Subtle differences in two non‐native congeneric beach grasses significantly affect their colonization,spread, and impact

Comparisons of congeneric species have provided unique insights into invasion ecology. Most often, non‐native species are compared to native ones to look for traits predicting invasion success. In this study, we examine a different facet of congeneric comparisons in which both species are non‐native. Ecological variability among non‐native congeners might 1) lead to the inhibition or facilitation of either species’ ability to colonize and spread, 2) result in larger cumulative impacts due to synergies between species, and 3) depend on the physical context of the invaded habitat. To explore these possibilities, we studied the distribution and abundance of two non‐native beach grasses: European beach grass Ammophila arenaria and American beach grass Ammophila breviligulata, their interaction with one another, and their biotic and physical impacts on dune ecosystems of the Pacific coast of North America. We found that over a two‐decade period, A. breviligulata has increased its dominance over A. arenaria on dunes where it was originally planted in 1935 and has actively spread to new sites formerly dominated by A. arenaria. Our results also show that dune plant species richness was lower at A. breviligulata sites, although there was an increase in the native beach grass Elymus mollis. More significantly, we found that the two grass species are associated with significantly different foredune shapes that are likely controlled by a combination of variability in sand supply along the coast and subtle differences in the congeners’ morphology and growth form. These differences have significant implications for the coastal protection services of dunes to humans and the conservation of native species. They provide a cautionary tale on the impacts of introducing novel species based purely on analogy with closely related species.     

Does salt stress constrain spatial distribution of dune building grasses Ammophila arenaria and Elytrichia juncea on the beach?

Rising sea levels threaten coastal safety by increasing the risk of flooding. Coastal dunes provide a natural form of coastal protection. Understanding drivers that constrain early development of dunes is necessary to assess whether dune development may keep pace with sea‐level rise. In this study, we explored to what extent salt stress experienced by dune building plant species constrains their spatial distribution at the Dutch sandy coast. We conducted a field transplantation experiment and a glasshouse experiment with two dune building grasses Ammophila arenaria and Elytrigia juncea. In the field, we measured salinity and monitored growth of transplanted grasses in four vegetation zones: (I) nonvegetated beach, (II) E. juncea occurring, (III) both species co‐occurring, and (IV) A. arenaria dominant. In the glasshouse, we subjected the two species to six soil salinity treatments, with and without salt spray. We monitored biomass, photosynthesis, leaf sodium, and nutrient concentrations over a growing season. The vegetation zones were weakly associated with summer soil salinity; zone I and II were significantly more saline than zones III and IV. Ammophila arenaria performed equally (zone II) or better (zones III, IV) than E. juncea, suggesting soil salinity did not limit species performance. Both species showed severe winter mortality. In the glasshouse, A. arenaria biomass decreased linearly with soil salinity, presumably as a result of osmotic stress. Elytrigia juncea showed a nonlinear response to soil salinity with an optimum at 0.75% soil salinity. Our findings suggest that soil salinity stress either takes place in winter, or that development of vegetated dunes is less sensitive to soil salinity than hitherto expected.     

Invasive Congeners Differ in Successional Impacts across Space and Time

Invasive species can alter the succession of ecological communities because they are often adapted to the disturbed conditions that initiate succession. The extent to which this occurs may depend on how widely they are distributed across environmental gradients and how long they persist over the course of succession. We focus on plant communities of the USA Pacific Northwest coastal dunes, where disturbance is characterized by changes in sediment supply, and the plant community is dominated by two introduced grasses – the long-established Ammophila arenaria and the currently invading A. breviligulata. Previous studies showed that A. breviligulata has replaced A. arenaria and reduced community diversity. We hypothesize that this is largely due to A. breviligulata occupying a wider distribution across spatial environmental gradients and persisting in later-successional habitat than A. arenaria. We used multi-decadal chronosequences and a resurvey study spanning 2 decades to characterize distributions of both species across space and time, and investigated how these distributions were associated with changes in the plant community. The invading A. breviligulata persisted longer and occupied a wider spatial distribution across the dune, and this corresponded with a reduction in plant species richness and native cover. Furthermore, backdunes previously dominated by A. arenaria switched to being dominated by A. breviligulata, forest, or developed land over a 23-yr period. Ammophila breviligulata likely invades by displacing A. arenaria, and reduces plant diversity by maintaining its dominance into later successional backdunes. Our results suggest distinct roles in succession, with A. arenaria playing a more classically facilitative role and A. breviligulata a more inhibitory role. Differential abilities of closely-related invasive species to persist through time and occupy heterogeneous environments allows for distinct impacts on communities during succession.     

Dominant regions and drivers of the variability of the global land carbon sink across timescales

Net biome productivity (NBP) dominates the observed large variation of atmospheric CO₂ annual increase over the last five decades. However, the dominant regions controlling inter‐annual to multi‐decadal variability of global NBP are still controversial (semi‐arid regions vs. temperate or tropical forests). By developing a theory for partitioning the variance of NBP into the contributions of net primary production (NPP) and heterotrophic respiration (R_h) at different timescales, and using both observation‐based atmospheric CO₂ inversion product and the outputs of 10 process‐based terrestrial ecosystem models forced by 110‐year observational climate, we tried to reconcile the controversy by showing that semi‐arid lands dominate the variability of global NBP at inter‐annual (<10 years) and tropical forests dominate at multi‐decadal scales (>30 years). Results further indicate that global NBP variability is dominated by the NPP component at inter‐annual timescales, and is progressively controlled by R_h with increasing timescale. Multi‐decadal NBP variations of tropical rainforests are modulated by the Pacific Decadal Oscillation (PDO) through its significant influences on both temperature and precipitation. This study calls for long‐term observations for the decadal or longer fluctuations in carbon fluxes to gain insights on the future evolution of global NBP, particularly in the tropical forests that dominate the decadal variability of land carbon uptake and are more effective for climate mitigation.     

Indigenous and Ammophila arenaria-dominated dune vegetation on the South African Cape coast

Abstract. Communities formed by the potentially invasive European Ammophila arenaria (marram grass) are compared with those dominated by indigenous dune plant species in coastal dune systems. Sampling of communities was carried out along the Cape coast for species richness, species diversity, importance values and species associations. The influence of soil and other environmental factors on vegetation were also compared. While species richness values in A. arenaria communities appear similar to those of indigenous dune plant communities, diversity indices are significantly lower. However, on the basis of importance values of individual species, A. arenaria does not show extreme dominance to the exclusion of other species, as it does on the North American Pacific coast, where it has also been introduced. Because of its growth in dense tufts, A. arenaria is accompanied mostly by small chamaephytes and therophytes, while indigenous stands support more phanerophytes. Moreover, A. arenaria forms weaker species associations than dominant indigenous dune plant species. The alien status of A. arenaria in South Africa is confirmed by applying classification and ordination analyses which failed to differentiate A. arenaria communities according to their geographical origin as achieved with indigenous communities. This may be attributed to the lack of vigorous indigenous plants in A. arenaria communities, which accounts for the low variety in species composition of A. arenaria communities along the coastline. With regard to environmental factors, A. arenaria communities were observed to be less sensitive to extrinsic factors, such as climate, than indigenous dune plant communities. Results confirm that A. arenaria is an alien plant species in South Africa, but do not imply its invasiveness in the present or near future.     

Seeking shelter from the storm: Conservation and management of imperiled species in a changing climate

Climate change is anticipated to exacerbate the extinction risk of species whose persistence is already compromised by habitat loss, invasive species, disease, or other stressors. In coastal areas of the southeastern United States (USA), many imperiled vertebrates are vulnerable to hurricanes, which climate models predict to become more severe in the 21st century. Despite this escalating threat, explicit adaptation strategies that address hurricane threats, in particular, and climate change more generally, are largely underrepresented in recovery planning and implementation. We provide a basis for stronger emphasis on strategic planning for imperiled species facing the increasing threat of catastrophic hurricanes. Our reasoning comes from observations of short‐term environmental and biological impacts of Hurricane Michael, which impacted the Gulf Coast of the southeastern USA in October 2018. During this storm, St. Marks National Wildlife Refuge, located along the northern Gulf of Mexico's coast in the panhandle region of Florida, received storm surge that was 3.0–3.6 m (NAVD88) above sea level. Storm surge pushed sea water into some ephemeral freshwater ponds used for breeding by the federally threatened frosted flatwoods salamander (Ambystoma cingulatum). After the storm, specific conductance across all ponds measured varied from 80 to 23,100 µS/cm, compared to 75 to 445 µS/cm in spring 2018. For 17 overwashed wetlands that were measured in both spring and fall 2018, posthurricane conductance observations were, on average, more than 90 times higher than in the previous spring, setting the stage for varying population responses across this coastal landscape. Importantly, we found live individual flatwoods salamanders at both overwashed and non‐overwashed sites, although we cannot yet assess the demographic consequences of this storm. We outline actions that could be incorporated into climate adaptation strategies and recovery planning for imperiled species, like A. cingulatum, that are associated with freshwater coastal wetlands in hurricane‐prone regions.     

Colonization of the root zone ofAmmophila arenaria by harmful soil organisms

The role of harmful soil organisms in the degeneration ofAmmophila arenaria at coastal foredunes was examined by the growing of seedlings ofA. arenaria in soil samples collected from its root zone. Three sites, each representing a successive stage in foredune succession were examined: (1) a highly mobile dune (sand accretion of 80 cm year⁻¹) with vigorousA. arenaria, colonizing only the upper 30-cm of the annually deposited layer of sand, (2) a mobile dune with vigorousA. arenaria (sand accretion of 22 cm year⁻¹) and a 1-metre soil profile completely colonized by roots and (3) a stable dune (no sand accretion) with degeneratedA. arenaria and young roots mainly present in the upper 0–10 cm. In the upper part of the highly mobile site, the presence of harmful soil organisms was confined to the root layers and at the mobile site for all depth layers a significant growth reduction ofA. arenaria was observed due to the activity of harmful soil organisms. At the stable site, however, growth had only been reduced in some of the depth layers. At all sites newly formed roots ofA. arenaria had been colonized by harmful soil organisms within one year. If present in sand prior to root growth harmful soil organisms reduced root length and root hair formation severely and they enhanced branching of the roots. It is concluded that harmful soil organisms initiate degeneration ofA. arenaria in stable dunes by attack of the root system, which makes the plants suffer from abiotic stress.     

The effect of gap dynamics on the size and spatial structure of Solidago sempervirens on primary coastal dunes

Abstract. The role of gap dynamics in shaping the spatial and size distributions of the gap specialist Solidago sempervirens (seaside goldenrod) were examined on the primary dune grassland (Ammophila breviligulata, American beachgrass) of an Atlantic barrier island. Results suggest that coastal dune blowouts are patchy disturbances that allow S. sempervirens to temporarily establish and disperse seed. Natural measurements and clipping experiments demonstrated that A. breviligulata competitively dominated S. sempervirens in undisturbed grasslands. Coastal dune blow-outs reversed this dominance by temporarily burying A. breviligulata and increasing densities and above-ground biomass of S. sempervirens. Examination of size/density distributions of S. sempervirens suggested that blow-outs were seed sources but not regeneration sites. Regeneration was found primarily in neighboring transition and undisturbed grassland areas. Comparison of lateral growth by A. breviligulata on blow-out margins suggested that disturbances were closed by lateral encroachment from gap edges. After blow-out recovery, greater densities and skewed size distributions of S. sempervirens could still be detected for a number of years after blow-out recovery.     

If you build it,will they come? Use of restored dunes by beach mice

Restoration of coastal habitat fragmented, degraded, or destroyed by development and climate‐related processes such as sea level rise and storm surge usually involves planting native plants to restore habitat structure, but whether and how restored areas benefit taxa other than plants is rarely reported. Installing restoration plantings is one method used to build habitat such as beach dunes where dunes have been lost, potentially creating habitat for dune‐dependent species. We compared use of natural vegetated dunes, open sand gaps, and restoration plantings (habitat treatment) by Perdido Key beach mice (Peromyscus polionotus trissyllepsis) over 3 years using tracking tubes to assess the value of restoration plantings for beach mice. Tubes were monitored in two seasons (early and mid‐summer), and under new and full moon conditions. Mice used restoration plantings less than natural vegetated dunes but more than open sand gaps, which suggests restoration plantings may facilitate movement of mice across fragmented areas. Both season and moon phase influenced the effect of habitat treatment, interactions which may be attributable to perceived risk associated with movement under a combination of different conditions of ambient light, vegetation cover, and habitat novelty. Our results show restoration plantings provide habitat for movement and foraging, and may ameliorate some consequences of sea level rise and storms for beach mice and potentially other dune‐dependent species into the future.     

A casualty of climate change? Loss of freshwater forest islands on Florida's Gulf Coast

Sea level rise elicits short‐ and long‐term changes in coastal plant communities by altering the physical conditions that affect ecosystem processes and species distributions. While the effects of sea level rise on salt marshes and mangroves are well studied, we focus on its effects on coastal islands of freshwater forest in Florida's Big Bend region, extending a dataset initiated in 1992. In 2014–2015, we evaluated tree survival, regeneration, and understory composition in 13 previously established plots located along a tidal creek; 10 plots are on forest islands surrounded by salt marsh, and three are in continuous forest. Earlier studies found that salt stress from increased tidal flooding prevented tree regeneration in frequently flooded forest islands. Between 1992 and 2014, tidal flooding of forest islands increased by 22%–117%, corresponding with declines in tree species richness, regeneration, and survival of the dominant tree species, Sabal palmetto (cabbage palm) and Juniperus virginiana (southern red cedar). Rates of S. palmetto and J. virginiana mortality increased nonlinearly over time on the six most frequently flooded islands, while salt marsh herbs and shrubs replaced forest understory vegetation along a tidal flooding gradient. Frequencies of tidal flooding, rates of tree mortality, and understory composition in continuous forest stands remained relatively stable, but tree regeneration substantially declined. Long‐term trends identified in this study demonstrate the effect of sea level rise on spatial and temporal community reassembly trajectories that are dynamically re‐shaping the unique coastal landscape of the Big Bend.     

Non‐native mangroves support carbon storage,sediment carbon burial,and accretion of coastal ecosystems

Mangrove forests play an important role in climate change adaptation and mitigation by maintaining coastline elevations relative to sea level rise, protecting coastal infrastructure from storm damage, and storing substantial quantities of carbon (C) in live and detrital pools. Determining the efficacy of mangroves in achieving climate goals can be complicated by difficulty in quantifying C inputs (i.e., differentiating newer inputs from younger trees from older residual C pools), and mitigation assessments rarely consider potential offsets to CO₂ storage by methane (CH₄) production in mangrove sediments. The establishment of non‐native Rhizophora mangle along Hawaiian coastlines over the last century offers an opportunity to examine the role mangroves play in climate mitigation and adaptation both globally and locally as novel ecosystems. We quantified total ecosystem C storage, sedimentation, accretion, sediment organic C burial and CH₄ emissions from ~70 year old R. mangle stands and adjacent uninvaded mudflats. Ecosystem C stocks of mangrove stands exceeded mudflats by 434 ± 33 Mg C/ha, and mangrove establishment increased average coastal accretion by 460%. Sediment organic C burial increased 10‐fold (to 4.5 Mg C ha^?1 year^?1), double the global mean for old growth mangrove forests, suggesting that C accumulation from younger trees may occur faster than previously thought, with implications for mangrove restoration. Simulations indicate that increased CH₄ emissions from sediments offset ecosystem CO₂ storage by only 2%–4%, equivalent to 30–60 Mg CO₂‐eq/ha over mangrove lifetime (100 year sustained global warming potential). Results highlight the importance of mangroves as novel systems that can rapidly accumulate C, have a net positive atmospheric greenhouse gas removal effect, and support shoreline accretion rates that outpace current sea level rise. Sequestration potential of novel mangrove forests should be taken into account when considering their removal or management, especially in the context of climate mitigation goals.     

Historical changes in the distributions of invasive and endemic marine invertebrates are contrary to global warming predictions: the effects of decadal climate oscillations

Aim We tested whether a hybrid zone that has formed between an endemic and an invasive species of marine mussel has shifted poleward as expected under a general hypothesis of global warming or has responded instead to decadal climate oscillations. Location We sampled 15 locations on the coast of California, USA, that span the distributions of the two species of marine mussels and their hybrids. Methods Mussels were sampled in 2005–08 and analysed at three nuclear gene loci using methods identical to those used in a study a decade earlier in order to document the genetic architecture of this system. Change in the system was determined by comparing the frequency of species‐specific alleles and multi‐locus genotypes over the intervening decade. Climate variation over the same period was examined by comparing the Pacific Decadal Oscillation (PDO), El Niño/Southern Oscillation (ENSO), upwelling indices and sea surface temperature (SST) during and prior to the study period. Results Contrary to the general expectations of global warming we show that the highly invasive warm‐water mussel Mytilus galloprovincialis and the hybrid zone formed with the endemic species Mytilus trossulus has rapidly contracted southwards. Mytilus galloprovincialis declined in abundance over the northern third of its geographic range (c. 540 km) and has become rare or absent across the northern 200 km of the range it previously colonized during its initial invasion. The distribution of the native species M. trossulus has remained unchanged over the same time period. Main conclusions The large‐scale range shift in the warm‐water invasive species M. galloprovincialis and the hybrid zone it forms with M. trossulus has been exceptionally rapid and is in the opposite direction to that predicted by the global warming hypotheses. This shift, however, is consistent with decadal climate variation associated with the ENSO and the PDO. Since the biogeography of this system was first described in 1999, the PDO has shifted from a warm phase, dominated by frequent and large El Niño events, to a cold‐phase period, with minimal ENSO activity. Thus recent decadal climate variation can oppose global trends in average temperature and this study illustrates the need to integrate the effects of climate change across multiple time‐scales.     

Extremely low genetic diversity across mangrove taxa reflects past sea level changes and hints at poor future responses

The projected increases in sea levels are expected to affect coastal ecosystems. Tropical communities, anchored by mangrove trees and having experienced frequent past sea level changes, appear to be vibrant at present. However, any optimism about the resilience of these ecosystems is premature because the impact of past climate events may not be reflected in the current abundance. To assess the impact of historical sea level changes, we conducted an extensive genetic diversity survey on the Indo‐Malayan coast, a hotspot with a large global mangrove distribution. A survey of 26 populations in six species reveals extremely low genome‐wide nucleotide diversity and hence very small effective population sizes (N_e) in all populations. Whole‐genome sequencing of three mangrove species further shows the decline in N_e to be strongly associated with the speed of past changes in sea level. We also used a recent series of flooding events in Yalong Bay, southern China, to test the robustness of mangroves to sea level changes in relation to their genetic diversity. The events resulted in the death of half of the mangrove trees in this area. Significantly, less genetically diverse mangrove species suffered much greater destruction. The dieback was accompanied by a drastic reduction in local invertebrate biodiversity. We thus predict that tropical coastal communities will be seriously endangered as the global sea level rises. Well‐planned coastal development near mangrove forests will be essential to avert this crisis.     

Impacts of simulated climate change and fungal symbionts on survival and growth of a foundation species in sand dunes

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.     

Feedback of coastal marshes to climate change: Long‐term phenological shifts

Coastal marshes are important carbon sinks facing serious threats from climatic stressors. Current research reveals that the growth of individual marsh plants is susceptible to a changing climate, but the responses of different marsh systems at a landscape scale are less clear. Here, we document the multi‐decadal changes in the phenology and the area of the extensive coastal marshes in Louisiana, USA, a representative of coastal ecosystems around the world that currently experiencing sea‐level rise, temperature warming, and atmospheric CO₂ increase. The phenological records are constructed using the longest continuous satellite‐based record of the Earth's ecosystems, the Landsat data, and an advanced modeling technique, the nonlinear mixed model. We find that the length of the growing seasons of the intermediate and brackish marshes increased concomitantly with the atmospheric CO₂ concentration over the last 30 years, and predict that such changes will continue and accelerate in the future. These phenological changes suggest a potential increase in CO₂ uptake and thus a negative feedback mechanism to climate change. The areas of the freshwater and intermediate marshes were stable over the period studied, but the areas of the brackish and saline marshes decreased substantially, suggesting ecosystem instability and carbon storage loss under the anticipated sea‐level rise. The marshes' phenological shifts portend their potentially critical role in climate mitigation, and the different responses among systems shed light on the underlying mechanisms of such changes.     

Fluctuating deposition of ocean water drives plant function on coastal sand dunes

Sea‐level rise will alter the hydrology of terrestrial coastal ecosystems. As such, it becomes increasingly important to decipher the present role of ocean water in coastal ecosystems in order to assess the coming effects of sea‐level rise scenarios. Sand dunes occur at the interface of land and sea. Traditionally, they are conceived as freshwater environments with rain and ground water as the only water sources available to vegetation. This study investigates the possibility of ocean water influx to dune soils and its effect on the physiology of sand dune vegetation. Stable isotopes are used to trace the path of ocean water from the soil to the vegetation. Soil salinity, water content and δ¹⁸O values are measured concurrently with stem water and leaf tissue of eight species during the wet and dry season and from areas proximal and distal to the ocean. Our results indicate the dune ecosystem is a mixed freshwater and marine water system characterized by oceanic influence on dune hydrology that is spatially heterogeneous and fluctuates temporally. Ocean water influx to soil occurs via salt spray in areas 5–12 m from the ocean during dry season. Accordingly, vegetation nearest to the sea demonstrate a plastic response to ocean water deposition including elevated integrated water use efficiency (δ¹³C_leaf) and uptake of ocean water that comprised up to 52% of xylem water. We suggest physiological plasticity in response to periodic ocean water influx may be a functional characteristic common to species on the leading edge of diverse coastal habitats and an important feature that should be included in modeling coastal ecosystems. Rising sea level would likely cause a repercussive landward shift of dune species in response to encroaching maritime influences. However, human development would restrict this process, potentially causing the demise of dune systems and the protection from land erosion they provide.     

Mycorrhizae and succession in plantings of beachgrass in sand dunes

A survey of arbuscular mycorrhizal fungi (AMF), arbuscular mycorrhizae (AM), and hyphal networks of AMF was carried out in sand dune sites of different successional stages in the Province Lands Area of Cape Cod National Seashore, Massachusetts. The study focused on large-scale plantings (each of 12–20 ha) of American beachgrass (Ammophila breviligulata) aged 0–7 yr and five adjacent natural dune areas. Sample sites ranged in vegetative cover from barren to forested. Spores of 17 species of AMF were recovered from the dunes. Over the successional sequence, there were increases in the richness and spore populations of the AMF community, the extent of colonization of A. breviligulata roots, and the mycorrhizal inoculum potential of the soil. Unvegetated sites lacked propagules of AMF, but roots of planted culms of A. breviligulata (which carried propagules of AMF) became mycorrhizal in <1 yr after planting. Spores were recovered from previously AMF-free sites that had been planted with beachgrass for 47 wk, and five species of AMF sporulated in sites <6 yr old. Significant hyphal networks were not present in any of the planted areas (<6 yr old at the time of sampling), but did occur in natural areas. The rate of invasion of areas planted to A. breviligulata by later successional plant species may in part depend upon the establishment of a vigorous network of hyphae of AMF in a site.     

Ecological Assessment of Dune Restorations in the Great Lakes Region

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.     

Optimal hurricane overwash thickness for maximizing marsh resilience to sea level rise

The interplay between storms and sea level rise, and between ecology and sediment transport governs the behavior of rapidly evolving coastal ecosystems such as marshes and barrier islands. Sediment deposition during hurricanes is thought to increase the resilience of salt marshes to sea level rise by increasing soil elevation and vegetation productivity. We use mesocosms to simulate burial of Spartina alterniflora during hurricane‐induced overwash events of various thickness (0–60 cm), and find that adventitious root growth within the overwash sediment layer increases total biomass by up to 120%. In contrast to most previous work illustrating a simple positive relationship between burial depth and vegetation productivity, our work reveals an optimum burial depth (5–10 cm) beyond which burial leads to plant mortality. The optimum burial depth increases with flooding frequency, indicating that storm deposition ameliorates flooding stress, and that its impact on productivity will become more important under accelerated sea level rise. Our results suggest that frequent, low magnitude storm events associated with naturally migrating islands may increase the resilience of marshes to sea level rise, and in turn, slow island migration rates. Synthesis: We find that burial deeper than the optimum results in reduced growth or mortality of marsh vegetation, which suggests that future increases in overwash thickness associated with more intense storms and artificial heightening of dunes could lead to less resilient marshes.     

Photosynthesis and water relations of four oak species: impact of flooding and salinity

 As global climate changes, sea level rise and increased frequency of hurricanes will expose coastal forests to increased flooding and salinity. Quercus species are frequently dominant in these forest, yet little is known about their salinity tolerance, especially in combination with flooding. In this study, 1-year-old seedlings of Quercus lyrata Walt. (overcup oak), Q. michauxii Nutt. (swamp chestnut oak), Q. nigra L. (water oak), and Q. nuttallii Palmer (Nuttall oak) were chronically (simulating sea level rise) and acutely (simulating hurricane storm surge) exposed to increased flooding and salinity, individually and in combination. The four species demonstrated two response patterns of photosynthesis (A), conductance, and leaf water potential, apparently related to their relative flood tolerance. In Q. lyrata, Q. nuttallii, and Q. nigra (moderately flood-tolerant), A was not immediately reduced after the initiation of the freshwater flooding, but was reduced as the duration of flooding increased. In the second pattern, demonstrated by the weakly flood-tolerant Q. michauxii, A was immediately reduced by freshwater flooding with an increasing impact over time. Watering with 2 parts per thousand (ppt) saline water did not consistently reduce A, but flooding with 2 ppt reduced A of all species, similar to the response with freshwater flooding. Photosynthesis of all species was reduced by 6 ppt watering or flooding, with the latter treatment killing all species within 8 weeks. When acutely exposed to 30 ppt salinity, A was quickly and severely reduced regardless of whether the seedlings were watered or flooded. Acutely flooded seedlings exposed to high salinity died within 2 weeks, but seedlings watered with 30 ppt saline water recovered and A was not reduced the following spring. As saline flooding of coastal areas increases due to sea level rise, photosynthesis of these species will be differentially affected based primarily on their flood tolerance. This suggests that increased flooding associated with sea level rise will impact these tree species to a greater extent than small increases in soil salinity. High salinity accompanying storm surges will be very harmful to all of these species. Received: 20 October 1997 / Accepted: 2 December 1998