Ammophila Invasion Ecology and Dune Restoration on the West Coast of North America

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.

Biomorphogenic Feedbacks and the Spatial Organization of a Dominant Grass Steer Dune Development

Nature-based solutions to mitigate the impact of future climate change depend on restoring biological diversity and natural processes. Coastal foredunes represent the most important natural flood barriers along coastlines worldwide, but their area has been squeezed dramatically because of a continuing urbanization of coastlines, especially in Europe. Dune development is steered by the development of vegetation in interaction with sand fluxes from the beach. Marram grass (Calamagrostis arenaria, formerly Ammophila arenaria) is the main dune building species along most European coasts, but also in other continents where the species was introduced. Engineering of coastal dunes, for instance by building dunes in front of dikes, needs to be based on a solid understanding of the species’ interactions with the environment. Only quantitative approaches enable the further development of mechanistic models and coastal management strategies that encapsulate these biomorphogenic interactions. We here provide a quantitative review of the main biotic and physical interactions that affect marram grass performance, their interactions with sand fluxes and how they eventually shape dune development. Our review highlights that the species’ spatial organization is central to dune development. We further demonstrate this importance by means of remote sensing and a mechanistic model and provide an outlook for further research on the use of coastal dunes as a nature-based solution for coastal protection.

Biomorphogenic feedbacks and the spatial organisation of a dominant grass steer dune development

Nature-based solutions to mitigate the impact of future climate change depend on restoring biological diversity and natural processes. Coastal foredunes represent the most important natural flood barriers along coastlines worldwide, but their area has been squeezed dramatically because of a continuing urbanisation of coastlines, especially in Europe. Dune development is steered by the development of vegetation in interaction with sand fluxes from the beach. Marram grass (Calamagrostis arenaria, formerly Ammophila arenaria) is the main dune building species along most European coasts, but also in other continents where the species was introduced. Engineering of coastal dunes, for instance by building dunes in front of dikes, needs to be based on a solid understanding of the species' interactions with the environment. Only quantitative approaches enable the further development of mechanistic models and coastal management strategies that encapsulate these biomorphogenic interactions. We here provide a quantitative review of the main biotic and physical interactions that affect marram grass performance, their interactions with sand fluxes and how they eventually shape dune development. Our review highlights that the species spatial organisation is central to dune development. We further demonstrate this importance by means of remote sensing and a mechanistic model and provide an outlook for further research on the use of coastal dunes as a nature-based solution for coastal protection.

The Adaptive Power of Ammophila arenaria: Biomimetic Study, Systematic Observation, Parametric Design, and Experimental Tests with Bimetal

The aim of our study was to apply a biomimetic approach, inspired by the Ammophila arenaria. This organism possesses a reversible leaf opening and closing mechanism that responds to water and salt stress (hydronastic movement). We adopted a problem-based biomimetic methodology in three stages: (i) two observation studies; (ii) how to abstract and develop a parametric model to simulate the leaf movement; and (iii) experiments with bimetal, a smart material that curls up when heated. We added creases to the bimetal active layer in analogy to the position of bulliform cells. These cells determine the leaf-closing pattern. The experiments demonstrated that creases influence and can change the direction of the bimetal natural movement. Thus, it is possible to replicate the Ammophila arenaria leaf-rolling mechanism in response to temperature variation and solar radiation in the bimetal. In future works, we will be able to propose responsive facade solutions based on these results.

Climate Change Alters The Interaction of Two Invasive Beachgrasses With Implications For Range Shifts And Coastal Dune Functions

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.

Scutellospora deformata (Scutellosporaceae), a new species of Gigasporales from the Mediterranean sand dunes of Spain

A new species S. deformata, that occurs in six locations of marine sand dunes along the eastern Mediterranean coast of Spain is described and illustrated from spores. In the field, the species occurred in the rhizosphere of Ammophila arenaria (Poaceae), Elymus farctus (Poaceae), Otanthus maritimus (Asteraceae), and Echinophora spinosa (Apiaceae). Morphological characters related with outer, middle and inner wall of the glomerospores as well as phylogenetic analysis (partial SSU, ITS1-5.8S region and the partial LSU nrDNA) support the hypothesis that the fungus is a new species of the Scutellosporaceae.

Invasion by Ammophila arenaria alters soil chemistry, leaving lasting legacy effects on restored coastal dunes in California

Many restoration projects rely on invasive plant removal to restore ecosystems. However, success of restoration efforts relying on invasives removal can be jeopardized, because, in addition to displacing native plants, invasives can also dramatically impact soils. Many studies have documented invasives’ effects on soil chemistry and microbiota. While European beachgrass (Ammophila arenaria (L.) Link) is a worldwide invasives problem in coastal dunes outside northern Europe, little attention has been paid to effects of this species on soil chemistry following invasion, even though it establishes persistent, dense monocultures. In our study, we evaluated effects of A. arenaria invasion on soil chemistry of coastal dunes at Point Reyes National Seashore (PRNS); persistence of effects following removal by mechanical or herbicide treatment (legacy effects); and effects of treatment independent of invasion. Dune restoration efforts at PRNS have met with mixed success, especially in herbicide-treated backdunes, where decomposition of dead A. arenaria has been greatly delayed. Based on results, invasion impacted 74% of 19 variables assessed, although there was a significant interaction in many cases with successional status (earlier vs later). Almost 60% of invasion effects persisted after restoration, with legacy effects prevalent in herbicide-treated backdunes where sand deposition from adjacent beaches could not mitigate effects as it could in herbicide-treated foredunes. Mechanical removal — or inversion of invaded surface soils with less-contaminated subsoils — resulted in fewer legacy effects, but more treatment effects, primarily in backdunes. Soil chemistry may decelerate decomposition of A. arenaria due to the limited nitrogen (N) available to enable microbial breakdown of the high carbon(C):N (70.8:1) material, but microbial factors probably play a more important role. Success of restoration at PRNS may not be fully realized until legacy effects are resolved through additional actions such as inoculation with healthy microbiomes or necromass reduction through controlled burning.

Coastal Observations: Rapid development and colonization of a UK sand dune system

Between the years 1200 and 1600, vast quantities of sand were brought inshore from offshore bars as a result of centuries of ferocious storms, to form a series of dune systems along the South Wales coastline. Today, as a result of many housing, leisure, and industrial developments only a few remnants exist. On one such remnant at Porthcawl, Wales, UK, became a caravan site in the 1930s, which was abandoned in 1993 for political reasons. Within 27 years a minimum of 120,000 m3 of sand was transported from the adjacent beach and formed dunes >4 m in height along a 400- m frontal edge that extended some 130 m inland, approximately a third of the site. Typical vegetation found along the frontal part of the system are Ammophila arenaria (marram), Agropyron junceiforme (sand couch grass) and Euphorbia maritimum (spurge). To the rear of the system, vegetation included Agrostis tenuis and stolonifera, (bent and creeping bent grass), Cirsium avense (creeping thistle), and Caluna vulgaris (heather). A 4-m-high and c. 3000m2 area of a vigorous stand of Hippophae rhamnoides (sea buckthorn) has also formed. The rapidity of dune formation and vegetation colonization is staggering.

Variation in Growth Response of Coastal Dune-Building Grass Species Ammophila Arenaria and Leymus Arenarius to Sand Burial

Ammophila arenaria and Leymus arenarius are dune-building grass species native to European seacoasts. The present study aimed to compare growth responses to the sand burial of A. arenaria and L. arenarius from coastal habitats of the Baltic Sea, when the intensity of sand accretion was relatively low under controlled conditions. Plants were grown from seeds collected from natural coastal habitats, transplanted into individual containers, buried in the sand at different depths in the rapid shoot elongation stage, and further cultivated (11 or 9 weeks) in an automated greenhouse. Burial in sand significantly stimulated the growth of shoots of A. arenaria, the effect was earlier at high burial intensities (46 and 60%) and was evident ten days after the start of treatment. Both shoot and root dry mass increased for plants buried at 13%; however, increased burial depth (37, 46 and 60%) resulted in a significant increase in root biomass. In comparison, shoot biomass decreased significantly at the highest burial intensity (60%). For L. arenarius, there was no direct dependence of shoot elongation rate on burial depth. There was a tendency for increased elongation growth and biomass allocation to leaf sheaths despite a decrease in total shoot mass. Most strikingly, root biomass decreased with sand burial in parallel with increased burial depth up to 21% intensity. In conclusion, although both grass species showed a positive shoot growth response to moderate sand burial intensity, differences in individual responses at the morphological and physiological level indicate the existence of different genetically based adaptation strategies.