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''Ammophila'' (synonymous with ''Psamma'' P. Beauv.) is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s consisting of two or three very similar species of
grasses Poaceae ( ), also called Gramineae ( ), is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos, the grasses of natural grassland and species cultivated in ...
. The genus name ''Ammophila'' originates from the Greek words ἄμμος (''ámmos''), meaning "sand", and φίλος (''philos''), meaning "friend". The
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often con ...
s for the grasses include marram grass, bent grass, and beachgrass. The grasses are found almost exclusively on the first line of
coast A coast (coastline, shoreline, seashore) is the land next to the sea or the line that forms the boundary between the land and the ocean or a lake. Coasts are influenced by the topography of the surrounding landscape and by aquatic erosion, su ...
al
dunes A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
. Their extensive system of creeping underground stems or
rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
s allows them to thrive under conditions of shifting sands and high winds, and helps stabilize the dunes and prevent
coastal erosion Coastal erosion is the loss or displacement of land, or the long-term removal of sediment and rocks along the coastline due to the action of Wind wave, waves, Ocean current, currents, tides, wind-driven water, waterborne ice, or other impacts ...
. ''Ammophila'' species are native to the coasts of the
North Atlantic Ocean The Atlantic Ocean is the second largest of the world's five oceanic divisions, with an area of about . It covers approximately 17% of Earth's surface and about 24% of its water surface area. During the Age of Discovery, it was known for se ...
where they are usually the dominant species on dunes. Their native range includes few inland regions, with the
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes spanning the Canada–United States border. The five lakes are Lake Superior, Superior, Lake Michigan, Michigan, Lake Huron, H ...
of North America being the main exception. The ''Ammophila'' grasses are widely known as examples of
xerophyte A xerophyte () is a species of plant that has adaptations to survive in an environment with little liquid water. Examples of xerophytes include cacti, pineapple and some gymnosperm plants. The morphology and physiology of xerophytes are adapted to ...
s, plants that can withstand dry conditions. Despite their occurrence on seacoasts, ''Ammophila'' grasses are not particularly tolerant of saline soils, although they can tolerate a salinity of about 15 g/L (1.5%), which makes them "moderate
halophyte A halophyte is a salt-tolerant plant that grows in soil or waters of high salinity, coming into contact with saline water through its roots or by salt spray, such as in saline semi-deserts, mangrove swamps, marshes and sloughs, and seashores. ...
s". ''Ammophila'' builds coastal dunes and thus stabilizes the sand. For that reason, the plants are seen as a useful means of reinforcing dunes around the world and have been introduced far from their native range. Alfred Wiedemann writes that ''
Ammophila arenaria ''Ammophila arenaria'' is a species of grass in the family Poaceae. It is known by the common names marram grass and European beachgrass. It is one of two species of the genus '' Ammophila''. It is native to the coastlines of Europe and North Afr ...
'', the European species, "has been introduced into virtually every British colonial settlement within its latitudinal tolerance range, including south-east and south-west Australia, New Zealand, South Africa, the Falkland Islands, and Norfolk Island, and has been reported from Argentina and Chile." ''Ammophila'' species were introduced in the late 19th century on the Pacific coast of North America as well, and massive, intentional plantings were continued at least through 1960. In virtually all of the locations where it has been introduced, ''Ammophila'' is now listed as invasive. ''Ammophila'' infestations adversely affect coastal groundbirds and endemic dune plants, such as the
western snowy plover The western snowy plover (''Anarhynchus nivosus nivosus'') is a small wader in the plover bird family. They are currently federally listed under the U.S. Endangered Species Act as Threatened. Human activity, habitat loss and predation are the bigg ...
, streaked horned lark (''Eremophila alpestris strigata''), and pink sand verbena (''Abronia umbellata'' subsp. ''breviflora''). In California, efforts to cull invasive ''Ammophila arenaria'' from
Morro Strand State Beach Morro Strand State Beach (formerly Atascadero State Beach) is a protected beach within California's state park system located in north Morro Bay, California, Morro Bay along California State Route 1. Morro Strand is a popular coastal frontag ...
began in 2000 and involved a combination of herbicide treatments and handpulling by a
California Conservation Corps The California Conservation Corps, CCC, is a department of the government of California, falling under the state Cabinet (government), cabinet-level California Resources Agency. The CCC is a voluntary work development program specifically for me ...
crew. That initiative produced a 60 percent mortality among the ''Ammophila'' after several months, but it was estimated that several additional rounds of treatment would be needed to fully remove the infestation, and the effort specifically focused on "relatively new infestations." Bulldozing has also been used as an effective mechanical treatment for ''Ammophila'' infestations in the Pacific Northwest, but research indicates that the intensity and frequency of bulldozing employed to remove ''Ammophila'' may also harm endemic dune plants and have negative impacts on dune geomorphology.


Species

Only two species seem incontrovertible: ''A. arenaria'' and ''A. breviligulata''. Two other species have been proposed, and are discussed below. * '' A. arenaria'' - European marram grass or European beachgrass. Native to coasts of
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
(north to
Iceland Iceland is a Nordic countries, Nordic island country between the Atlantic Ocean, North Atlantic and Arctic Oceans, on the Mid-Atlantic Ridge between North America and Europe. It is culturally and politically linked with Europe and is the regi ...
) and northwest
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
. Inflorescence to 25 cm long; broad. * ''A. baltica'' - Purple marram. ''A. baltica'' has now been identified as a
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two diff ...
between ''A. arenaria'' and ''
Calamagrostis epigejos ''Calamagrostis epigejos'', common names wood small-reed or bushgrass, is a species of grass in the family Poaceae which is native to Eurasia and Africa. It is found from average moisture locales to salt marsh and wet habitats.
''. The hybrid occurs in parts of northern Europe, mainly from the
Baltic Sea The Baltic Sea is an arm of the Atlantic Ocean that is enclosed by the countries of Denmark, Estonia, Finland, Germany, Latvia, Lithuania, Poland, Russia, Sweden, and the North European Plain, North and Central European Plain regions. It is the ...
west to eastern England, and is known as ''× Ammocalamagrostis baltica'' or ''× Calammophila baltica''. * '' A. breviligulata'' - American marram grass or American beachgrass. Native to coasts of eastern
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, including the shores of the
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes spanning the Canada–United States border. The five lakes are Lake Superior, Superior, Lake Michigan, Michigan, Lake Huron, H ...
. Inflorescence to 30 cm long; narrower than ''A. arenaria''. * ''A. champlainensis'' or ''A. breviligulata'' ssp. ''champlainensis'' - Champlain beachgrass. Native to the shores of
Lake Ontario Lake Ontario is one of the five Great Lakes of North America. It is bounded on the north, west, and southwest by the Canadian province of Ontario, and on the south and east by the U.S. state of New York (state), New York. The Canada–United Sta ...
and
Lake Champlain Lake Champlain ( ; , ) is a natural freshwater lake in North America. It mostly lies between the U.S. states of New York (state), New York and Vermont, but also extends north into the Canadian province of Quebec. The cities of Burlington, Ve ...
. Inflorescence to 22 cm long; very similar to ''A. breviligulata'', and no longer considered a distinct species by several authorities.


Ecology

In Europe, ''Ammophila arenaria'' has a coastal distribution and is the dominant species on
dunes A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
where it is responsible for stabilising and building the foredune by capturing blown sand and binding it together with the warp and weft of its tough, fibrous rhizome system. Marram grass is strongly associated with two coastal plant community types in the
British National Vegetation Classification __NOTOC__ The British National Vegetation Classification or NVC is a system of classifying natural habitat types in Great Britain according to the vegetation they contain. A large scientific meeting of ecologists, botanists, and other related ...
. In community SD6 (Mobile Dune) ''Ammophila'' is the dominant species. In the semi-fixed dunes (community SD7), where the quantity of blown sand is declining ''Ammophila'' becomes less competitive, and other species, notably ''
Festuca rubra ''Festuca rubra'' is a species of grass known by the common name red fescue, creeping red fescue or the rush-leaf fescue. It is widespread across much of the Northern Hemisphere and can tolerate many habitats and climates. It is best adapted to w ...
'' (red fescue) become prominent.


Uses

The ability of marram grass to grow on and bind sand makes it a useful plant in the stabilization of coastal
dune A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
s and artificial defences on sandy coasts. That usefulness was recognized in the late 18th century. On the
North Sea The North Sea lies between Great Britain, Denmark, Norway, Germany, the Netherlands, Belgium, and France. A sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the Norwegian Se ...
coast of
Jutland Jutland (; , ''Jyske Halvø'' or ''Cimbriske Halvø''; , ''Kimbrische Halbinsel'' or ''Jütische Halbinsel'') is a peninsula of Northern Europe that forms the continental portion of Denmark and part of northern Germany (Schleswig-Holstein). It ...
,
Denmark Denmark is a Nordic countries, Nordic country in Northern Europe. It is the metropole and most populous constituent of the Kingdom of Denmark,, . also known as the Danish Realm, a constitutionally unitary state that includes the Autonomous a ...
, marram grass was traditionally used for fuel, thatch, and cattle fodder after frost. Those uses led to sand drift and loss of arable land. Hence, legislation promoting dune stabilization came into force in 1779 and 1792, successively leading to a system of state-supported ''dune planters'' overlooked by ''dune bailiffs''. Marram grass was – and still is – propagated by root and shoot cuttings dug up locally and planted into the naked sand in periods of relatively calm and moist weather. Women from the village of
Newborough, Anglesey Newborough () is a village in the southern corner of the Isle of Anglesey in Wales. It is a part of the Bro Aberffraw electoral ward; to the northwest is Aberffraw, and to the northeast is Llanfairpwll. There is a church in the village, a prima ...
,
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
, once used marram grass in the manufacture of mats,
hay Hay is grass, legumes, or other herbaceous plants that have been cut and dried to be stored for use as animal fodder, either for large grazing animals raised as livestock, such as cattle, horses, goats, and sheep, or for smaller domesticate ...
stack covers and brushes for
whitewash Whitewash, calcimine, kalsomine, calsomine, asbestis or lime paint is a type of paint made from slaked lime ( calcium hydroxide, Ca(OH)2) or chalk (calcium carbonate, CaCO3), sometimes known as "whiting". Various other additives are sometimes ...
ing. Marram grass has been used for
thatch Thatching is the craft of building a roof with dry vegetation such as straw, Phragmites, water reed, Cyperaceae, sedge (''Cladium mariscus''), Juncus, rushes, Calluna, heather, or palm branches, layering the vegetation so as to shed water away fr ...
in many areas close to the sea in the
British Isles The British Isles are an archipelago in the Atlantic Ocean, North Atlantic Ocean off the north-western coast of continental Europe, consisting of the islands of Great Britain, Ireland, the Isle of Man, the Inner Hebrides, Inner and Outer Hebr ...
. The harvesting of marram grass for thatch was so widespread during the 17th century that it had the effect of destabilizing dunes, resulting in the burial of many villages, estates and farms. In 1695, the practice was banned by an Act of the Scottish Parliament:


Adaptations

Like other
xerophytes A xerophyte () is a species of plant that has adaptations to survive in an environment with little liquid water. Examples of xerophytes include cacti, pineapple and some gymnosperm plants. The morphology and physiology of xerophytes are adapted to ...
, marram grass is well-adapted to thrive in an otherwise harsh environment. The natural loss of water through
transpiration Transpiration is the process of water movement through a plant and its evaporation from aerial parts, such as leaves, stems and flowers. It is a passive process that requires no energy expense by the plant. Transpiration also cools plants, c ...
is not desirable in a very dry landscape, and marram grass has developed particular adaptations to help it deal with that. Sandy conditions drain water quickly, and very windy conditions will further increase rates of transpiration. Marram grass has a rolled leaf that creates a localized environment of water vapour concentration within the leaf and helps to prevent water loss. The
stomata In botany, a stoma (: stomata, from Greek ''στόμα'', "mouth"), also called a stomate (: stomates), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange between the internal air spa ...
sit in small pits within the curls of the structure, which makes them less likely to open and lose water. The folded leaves have hairs on the inside to slow or stop air movement, much like many other xerophytes. Although they are typically found on the outside of the plant, in marram grass they are also within the leaf as this has now become a structure with more volume. The slowing of air movement also reduces the amount of water vapour being lost. A waxy
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
on the leaf surface inhibits evaporation as well.


See also

*
List of Poaceae genera Poaceae, also known as the true grasses, is the fourth largest plant family in the world with around 12,000 species and roughly 800 genera. They contain, among others, the cereal crop species and other plants of economic importance, such as the b ...


References


External links

*
''California Conservation Corps
spearhead of ''Ammophila'' removal efforts in California.
Jepson Manual Treatment
{{Taxonbar, from=Q21130 Pooideae Flora of the Great Lakes region (North America) Poaceae genera