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''Ammophila breviligulata'' (American beachgrass or American marram grass) is a species of
grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns an ...
native to eastern
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, where it grows on sand
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, f ...
s along the
Atlantic Ocean The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe ...
and
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes in the mid-east region of North America that connect to the Atlantic Ocean via the Saint Lawrence River. There are five lakes ...
coasts. Beachgrass thrives under conditions of shifting sand, sand burial, and high winds; it is a dune-building grass that builds the first line of sand dunes along the coast. Beachgrass is less vigorous in stabilized sand, and is only infrequently found further inland than the coastal foredunes. On the Atlantic coastline of North America, ''Ammophila breviligulata'' has been observed as far south as North Carolina, and is often planted in dune restoration projects''. Ammophila breviligulata'' was introduced to the Pacific coast of North America in the 1930s. It is proving to be invasive, and is increasingly important to coastal ecology and development in
Oregon Oregon () is a U.S. state, state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington (state), Washington, while the Snake River delineates much of it ...
,
Washington Washington commonly refers to: * Washington (state), United States * Washington, D.C., the capital of the United States ** A metonym for the federal government of the United States ** Washington metropolitan area, the metropolitan area centered o ...
, and
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
.


Description

''Ammophila breviligulata'' is perennial grass which produces an extensive
rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...
system. The leaves have deeply furrowed upper surfaces and smooth undersides, and grow tall. The plant's
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
is a spike-like
panicle A panicle is a much-branched inflorescence. (softcover ). Some authors distinguish it from a compound spike inflorescence, by requiring that the flowers (and fruit) be pedicellate (having a single stem per flower). The branches of a panicle are of ...
that can reach long; The spikelet is single-flowered,
awn AWN may stand for: * Awn Access to Justice Network in Gaza Strip, Legal Aid Network operate in Gaza Strip, Palestine * Animation World Network, an online organization for animators * Avant Window Navigator, a dock-like bar that tracks open windows ...
less. The seed head appears in late July or August. The species name ''breviligulata'' derives from the Latin ''brevis'' ("short") and ''ligula'' ("tongue"), which refers to a feature of grass leaves called the
ligule A ligule (from "strap", variant of ''lingula'', from ''lingua'' "tongue") is a thin outgrowth at the junction of leaf and leafstalk of many grasses (Poaceae) and sedges. A ligule is also a strap-shaped extension of the corolla, such as that of a ...
. ''Ammophila breviligulata'' is quite similar in appearance and ecology to a second species of beachgrass, ''
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 ...
'' (European beachgrass). As Nick Page has summarized, "''Ammophila breviligulata'' is distinguished from ''A. arenaria'' by smaller ligules (1–3 mm versus 10–30 mm long in ''A. arenaria''), wider and less inrolled leaves, longer flower spike (25–35 cm versus 15–25 cm long in ''A. arenaria''), and scaly rather than puberulent leaf veins on the upper leaf surface."


Ecology

Both species of beachgrass are noted for their association with stable sand dunes, and observations in regions where they were introduced to coasts indicate that they build the first line of dunes on coasts (the foredunes). The plants spread rapidly – annually – through the sand by subsurface runners (
rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...
s), and can produce up to 100 stems per clump annually. They can tolerate burial in as much as of sand; sand burial stimulates the rhizomes to grow vertically, and is essential to plant vigor. The beachgrass species are also very good examples of
xerophyte A xerophyte (from Ancient Greek language, Greek ξηρός ''xeros'' 'dry' + φυτόν ''phuton'' 'plant') is a species of plant that has adaptations to survive in an environment with little liquid water, such as a desert such as the Sahara or pl ...
s, being able to thrive on arid to semi-arid beach dunes. The plant has several mechanisms for adapting to heat stress or wind. The long narrow leaves can roll or fold, and the rough upper leaf surface, which contains the gas exchange openings (stomata), can orient itself away from the wind. Beachgrass is a dominant species on foredunes, and is sometimes the only plant found there, but is unable to grow farther inland on stabilized dunes and
soil Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former te ...
. One reason for this that has been extensively studied by Wim van der Putten and his colleagues is based on the susceptibility of ''Ammophila'' species to soil pathogens such as
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s. The density of these pathogens is low in freshly deposited sand, but builds up in the roots of beachgrass on stabilized sand. While the details are complex, the result is that beachgrass is largely limited to the foredunes that it builds, which are immediately adjacent to the water's edge and where sand is removed and redeposited fairly frequently. Around the Great Lakes, it is limited to active dune habitats along the shores, and it is listed a threatened species in Minnesota where it reaches its most westerly natural range around Duluth.


Invasiveness

Starting in the 19th century, ''A. arenaria'' was introduced to the Pacific coast of North America to help control beach sands; it serves this role so well that it is now considered invasive not only along this coast (from
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
north to
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
), but along nearly every coast worldwide where it has been introduced. ''Ammophila arenaria'' built foredunes (the dunes closest to the beach) that are typically about high, which is much higher than the dunes associated with the native dune grass species ''
Leymus mollis ''Leymus mollis'' is a species of grass known by the common names American dune grass, American dune wild-rye, sea lyme-grass, strand-wheat,Higman, P. J. and M. R. Penskar. 1999Special plant abstract for ''Leymus mollis'' (American dune wild-rye) ...
''. In the 1930s, ''A. breviligulata'' was introduced to about on the
Clatsop Peninsula The Clatsop is a small tribe of Chinookan-speaking Native Americans in the Pacific Northwest of the United States. In the early 19th century they inhabited an area of the northwestern coast of present-day Oregon from the mouth of the Columbia ...
in
Oregon Oregon () is a U.S. state, state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington (state), Washington, while the Snake River delineates much of it ...
. Since then, ''A. breviligulata'' has gradually supplanted ''A. arenaria'' in Oregon and
Washington Washington commonly refers to: * Washington (state), United States * Washington, D.C., the capital of the United States ** A metonym for the federal government of the United States ** Washington metropolitan area, the metropolitan area centered o ...
; the precise reason for the relative success of ''A. breviligulata'' isn't known. The most recognizable aspect of the changeover between the beachgrass species is that the foredunes of ''A. breviligulata'' are shorter than the foredunes of ''A. arenaria''; foredunes of ''A. breviligulata'' are about high in this region. The diversity of other species that co-exist with mature stands of ''A. arenaria'' is comparable to the species diversity with ''A. breviligulata''. However, the lower foredune height for the latter means that coastal ecosystems that had adapted to the fairly tall foredunes of ''A. arenaria'' are again changing, especially in the regions just behind the foredune and sheltered by it. Similarly, buildings and roads that were constructed under the presumption of continuing protection by the tall ''A. arenaria'' foredunes may now be threatened by shifting sand and by increased exposure to wind and water in storms.


References

{{Taxonbar, from=Q2843697 Pooideae Grasses of the United States Grasses of Canada Flora of the Great Lakes region (North America) Flora of Eastern Canada Flora of the Eastern United States Plants described in 1920