Editing Ammophila (plant)

{{Short description|Genus of flowering plants}}
{{About-distinguish|grass|Ammophila (wasp)}}
{{automatic taxobox
|image = Helmgras_kijkduin_februari_2005.JPG
|image_caption = European marram grass <br />Dunes near [[The Hague]], February 2005
|taxon = Ammophila (plant)
|authority = [[Nicolaus Thomas Host|Host]]<ref>{{cite web
 |url          = http://www.ars-grin.gov/cgi-bin/npgs/html/splist.pl?532
 |title        = Species in GRIN for genus
 |publisher    = www.ars-grin.gov
 |access-date   = 2008-04-24
 |last         = 
 |archive-url  = https://web.archive.org/web/20081011172819/http://www.ars-grin.gov/cgi-bin/npgs/html/splist.pl?532
 |archive-date = 2008-10-11
 |url-status     = dead
}}</ref>
|subdivision_ranks = Species
|subdivision = *''[[Ammophila arenaria]]'' (L.) Link
* ''[[Ammophila breviligulata]]'' Fernald
* ''A. champlainensis'' Seymour
|range_map = Ammophila range map.jpg
|range_map_caption=The range of ''Ammophila'' in North America
|}}

'''''Ammophila''''' (synonymous with ''Psamma'' P. Beauv.) is a [[genus]] of [[flowering plant]]s consisting of two or three very similar species of [[Poaceae|grasses]]. The genus name ''Ammophila'' originates from the Greek words ἄμμος (''ámmos''), meaning "sand", and φίλος (''philos''), meaning "friend". The [[common name]]s for the grasses include '''marram grass''', '''bent grass''', and '''beachgrass'''.<ref>{{ITIS |id=40447 |taxon=''Ammophila'' |access-date=19 September 2008 }}</ref> The grasses are found almost exclusively on the first line of [[coast]]al [[dunes]]. Their extensive system of creeping underground stems or [[rhizome]]s allows them to thrive under conditions of shifting sands and high winds, and helps stabilize the dunes and prevent [[coastal erosion]].

''Ammophila'' species are native to the coasts of the [[Atlantic Ocean|North Atlantic Ocean]] where they are usually the dominant species on dunes. Their native range includes few inland regions, with the [[Great Lakes]] of North America being the main exception.<ref>{{cite book|last=Preston |first=C. D. |author2=Pearman, D. A.|author3= Dines, T. D. |year=2002 |title=New Atlas of the British and Irish Flora |publisher=Oxford University Press |isbn=978-0-19-851067-3}}</ref>

The ''Ammophila'' grasses are widely known as examples of [[xerophyte]]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&nbsp;g/L (1.5%), which makes them "moderate [[halophyte]]s".<ref>{{cite web |url=http://www.natureserve.org/explorer/servlet/NatureServe?searchName=Ammophila+arenaria |title=Comprehensive Report Species - Ammophila arenaria |publisher=[[NatureServe]] |access-date=2008-09-28 |archive-date=2009-07-25 |archive-url=https://web.archive.org/web/20090725065839/http://www.natureserve.org/explorer/servlet/NatureServe?searchName=Ammophila+arenaria |url-status=dead }}</ref>

''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]]'', 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."<ref>{{cite web|last=Wiedemann |first=Alfred |title=''Ammophila arenaria'' on the northwest coast of North America |date=1998-02-12 |url=http://www.ou.edu/cas/botany-micro/ben/ben183.html |work=Botanical Electronic News - BEN #183 |access-date=2008-09-19| archive-url= https://web.archive.org/web/20080927054149/http://www.ou.edu/cas/botany-micro/ben/ben183.html| archive-date= 27 September 2008 | url-status= live}}</ref> ''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 species|invasive]]. ''Ammophila'' infestations adversely affect coastal groundbirds and endemic dune plants, such as the [[western snowy plover]], [[horned lark|streaked horned lark (''Eremophila alpestris strigata'')]], and [[Abronia umbellata|pink sand verbena (''Abronia umbellata'' subsp. ''breviflora'')]].<ref name=":0">{{Cite journal |last=Zarnetske |first=Phoebe |last2=Seabloom |first2=Eric |last3=Hacker |first3=Sally |date=November 24, 2010 |title=Non-target effects of invasive species management: beachgrass, birds, and bulldozers in coastal dunes |url=https://esajournals.onlinelibrary.wiley.com/doi/10.1890/ES10-00101.1 |journal=Ecosphere |volume=1 |issue=5 |pages=1-20 |via=Wiley Online Library}}</ref> In California, efforts to cull invasive ''Ammophila arenaria'' from [[Morro Strand State Beach]] began in 2000 and involved a combination of herbicide treatments and handpulling by a [[California Conservation Corps]] 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."<ref>{{Cite web |title=Beach Grass Removal Project |url=https://www.parks.ca.gov/?page_id=21575 |access-date=November 7, 2024 |website=California Department of Parks and Recreation}}</ref> 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.<ref name=":0" />

== Species ==
Only two species seem incontrovertible: ''A.&nbsp;arenaria'' and ''A.&nbsp;breviligulata''. Two other species have been proposed, and are discussed below.
* '''''[[Ammophila arenaria|A. arenaria]]''''' - '''European marram grass''' or '''European beachgrass'''. Native to coasts of [[Europe]] (north to [[Iceland]]) and northwest [[Africa]]. Inflorescence to 25&nbsp;cm long; broad.
* '''''A. baltica''''' - '''Purple marram'''. ''A. baltica'' has now been identified as a [[Hybrid (biology)|hybrid]] between ''A. arenaria'' and ''[[Calamagrostis epigejos]]''. The hybrid occurs in parts of northern Europe, mainly from the [[Baltic Sea]] west to eastern England, and is known as ''× Ammocalamagrostis baltica'' or ''× Calammophila baltica''.<ref>{{cite web |title=Calamagrostis epigejos x Ammophila arenaria = X Calammophila baltica (Flüggé ex Schrad.) Brand |url=https://data.nbn.org.uk/Taxa/NHMSYS0001754369 |publisher=National Biodiversity Network (UK)}}</ref>
* '''''[[Ammophila breviligulata|A. breviligulata]]''''' - '''American marram grass''' or '''American beachgrass'''. Native to coasts of eastern [[North America]], including the shores of the [[Great Lakes]]. Inflorescence to 30&nbsp;cm long; narrower than ''A. arenaria''.
* '''''A. champlainensis''''' or '''''A.&nbsp;breviligulata''''' '''ssp.''' '''''champlainensis''''' - '''Champlain beachgrass'''. Native to the shores of [[Lake Ontario]] and [[Lake Champlain]]. Inflorescence to 22&nbsp;cm long; very similar to ''A. breviligulata'', and no longer considered a distinct species by several authorities.<ref>{{ITIS |id=185263 |taxon=''Ammophila champlainensis'' |access-date=6 September 2008 }}</ref><ref>The identification of Champlain beachgrass as a species that is distinct from ''A. breviligulata'' is still under investigation, and some authors consider Champlain beachgrass to be a subspecies ''A. breviligulata'' ssp. ''champlainensis'' (Seymour) P.J. Walker, C.A. Paris & Barrington ex Barkworth. Recent work on morphological differences between varying populations assigned to ''A. breviligulata'' and to ''A. champlainensis'' do not support the assignment of a distinct species for the latter specimens, despite significant differences. See {{cite journal |last=Delisle-Oldham |first=M. B. |author2=Oldham, M. J.|author3= Catling, P. M. |year=2008 |title=Taxonomic Recognition of Ammophila champlainensis and Morphological Variation in Northeastern North American Ammophila (Poaceae) |url=http://www.bioone.org/perlserv/?request=get-abstract&doi=10.3119%2F07-07.1&ct=1 |journal=Rhodora |volume=110 |issue=942 |pages=129–156 |access-date=2008-09-21 |doi=10.3119/07-07.1 |s2cid=86593875 }}</ref><ref>
{{cite web|title=Champlain Beachgrass Guide |publisher=New York Natural Heritage Program |url=http://www.acris.nynhp.org/guide.php?id=9716 |access-date=2008-09-27}}</ref>

== Ecology ==
{{unreferenced section|date=February 2016}}
In Europe, ''Ammophila arenaria'' has a coastal distribution and is the dominant species on [[sand dunes|dunes]] 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 [[Shingle, strandline and sand-dune communities in the British National Vegetation Classification system|coastal plant community types]] in the [[British National Vegetation Classification]]. 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]]'' (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]]s and artificial [[coastal management|defences]] on sandy coasts. That usefulness was recognized in the late 18th century.<ref>Viborg, E. (1788) Efterretning om Sandvexterne og deres Anvendelse til at dæmpe Sandflugten paa Vesterkanten af Jylland. [[Copenhagen|København]]. [https://archive.org/details/efterretningoms00vibogoog Full text (in Danish)](Subsequent [[German language|German]] translation: Beschreibung der Sandgewächse und ihrer Anwendung zur Hemmung des Flugsandes auf der Küste von Jütland etc. Viborg, Erich Aus dem Dänischen von J. Petersen. Kopenhagen, 1789)</ref><ref>Andresen, C. C. 1861. Om Klitformationen og Klittens Behandling og Bestyrelse. - P.G. Philipsens Forlag. [https://books.google.com/books?id=tFcDAAAAYAAJ&q=Andresen,+C.+C.+1861.+Om+Klitformationen. Full text (in Danish)]</ref> On the [[North Sea]] coast of [[Jutland]], [[Denmark]], 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]], [[Wales]], once used marram grass in the manufacture of mats, [[hay]]stack covers and brushes for [[whitewash]]ing.

Marram grass has been used for [[thatch]] in many areas close to the sea in the [[British Isles]]. 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:<ref>{{cite book|last=Gimingham |first=C. H. |year=1964 |chapter=Maritime and sub-maritime communities |title=The Vegetation of Scotland |editor=Burnett, J. H. |publisher=Oliver & Boyd |location=Edinburgh and London |pages=67–142}}</ref> {{quote|Considering that many lands, meadowes and pasturages lying on sea coasts have been ruined and overspread in many places in this kingdom by sand driven from adjacent sand hills ... His Majesty does strictly prohibit and discharge the pulling of bent, [[broom (shrub)|broom]] or [[juniper]] off the sand hills for hereafter.}}

== Adaptations ==
[[File:European Marram Grass (Ammophila arenaria) rolled leaf close-up.jpg|thumb|A single leaf of marram grass, showing the rolled leaf which reduces water loss]]

Like other [[xerophytes]], marram grass is well-adapted to thrive in an otherwise harsh environment. The natural loss of water through [[transpiration]] 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]] 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 [[plant cuticle|cuticle]] on the leaf surface inhibits evaporation as well.

== See also ==
* [[List of Poaceae genera]]

== References ==
{{Reflist|2}}

== External links ==
* {{Commons-inline}}
* [http://www.ccc.ca.gov ''California Conservation Corps], spearhead of ''Ammophila'' removal efforts in California.
* [http://ucjeps.berkeley.edu/cgi-bin/get_JM_treatment.pl?8738,8799 Jepson Manual Treatment]

{{Taxonbar|from=Q21130}}

[[Category:Pooideae]]
[[Category:Flora of the Great Lakes region (North America)]]
[[Category:Poaceae genera]]