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Distichlis spicata

Distichlis spicata (L.) Greene, commonly known as saltgrass, inland saltgrass, or seashore saltgrass, is a perennial, rhizomatous, sod-forming grass in the family Poaceae, renowned for its exceptional tolerance to saline, alkaline, and periodically flooded soils.^[1]^[2] It typically grows 6 to 35 inches (15–90 cm) tall with wiry, erect culms, flat or folded leaves arranged in two ranks, and extensive rhizomes and stolons that form dense mats up to 2,000–3,000 stems per square meter; the species is dioecious, with separate male and female plants producing yellowish to purplish spike-like inflorescences from May to October.^[3]^[4] Native to the Americas, D. spicata is widely distributed across North America—from southern Canada (provinces including Alberta, British Columbia, and Ontario) through the United States (in 43 states, including coastal regions from Maine to Florida and inland to the Great Basin) and Mexico—and extends southward into parts of South America.^[2]^[4] It thrives in diverse saline habitats such as coastal and inland salt marshes, alkali flats, playas, brackish meadows, desert washes, and disturbed sites with moist to intermittently flooded soils (pH 6.8–9.2, salinity 0.03%–5.4%), often serving as a pioneer species in early succession.^[3]^[4] Ecologically significant, D. spicata stabilizes shorelines, filters pollutants through its root systems, and supports biodiversity by providing forage, cover, and nesting sites for wildlife including waterfowl, fish, invertebrates, and the larvae of the wandering skipper butterfly (Panoquina panoquinoides errans); it also yields fair to good livestock forage (up to 9,081 kg/ha dry matter) and is used in reclamation and erosion control projects.^[3]^[4]^[5] Globally secure (G5 rank), the species faces localized threats from habitat alteration but recovers well from disturbances like fire and grazing due to its clonal growth and seed production.^[2]^[4]

Taxonomy

Classification

_Distichlis spicata belongs to the kingdom Plantae, phylum Tracheophyta, class Liliopsida, order Poales, family Poaceae, subfamily Chloridoideae, tribe Cynodonteae, genus Distichlis, and species D. spicata.^[6]^[7]^[8] The species has accumulated numerous synonyms over time, reflecting historical taxonomic revisions. Key synonyms include Distichlis stricta (Torr.) Rydb., Spartina spicata (L.) Kuntze, Uniola spicata L., Distichlis spicata var. divaricata Beetle, Distichlis spicata var. nana Beetle, Distichlis spicata var. stolonifera Beetle, Briza spicata (L.) Lam., Agropyron peruvianum (Lam.) Roem. & Schult., Brizopyrum americanum (L.) Link, and Poa thalassica Kunth.^[9]^[7]^[10] Two infraspecific varieties are commonly recognized within D. spicata: var. spicata and var. stricta. Var. spicata features shorter lemma awns measuring 0.2–0.5 mm and is primarily associated with coastal regions, while var. stricta has longer awns of 0.5–1.5 mm and occurs more frequently in inland saline areas; these differences are clinal and sometimes debated, leading some authorities to treat var. stricta as a distinct species (D. stricta).^[11]^[9]

Nomenclature and etymology

The scientific name Distichlis spicata derives from classical roots reflecting key morphological features of the plant. The genus name Distichlis originates from the Greek word distichos, meaning "two-ranked" or "in two rows," alluding to the characteristic arrangement of the leaves in two opposite ranks along the stem.^[12] The specific epithet spicata comes from the Latin spica, meaning "spike" or "ear of grain," referring to the dense, spike-like inflorescence structure.^[13] Common names for Distichlis spicata emphasize its ecological niche and form, with variations tied to regional habitats. The most widespread name, "saltgrass," stems from its prevalence in saline or alkaline soils, where it thrives as a dominant species in coastal marshes, inland playas, and desert fringes.^[6] Regional variants include "seashore saltgrass" for coastal populations, "inland saltgrass" for interior saline areas, "desert saltgrass" for arid southwestern occurrences, and "spike grass" highlighting the compact, spicate flower heads.^[3] The nomenclatural history of Distichlis spicata began with its original description by Carl Linnaeus as Uniola spicata in the 1753 edition of Species Plantarum, based on specimens from saline environments in the Americas.^[14] In 1887, American botanist Edward Lee Greene transferred the species to the genus Distichlis, establishing the current binomial in the Bulletin of the California Academy of Sciences, recognizing distinct generic traits such as the two-ranked leaves and dioecious nature.^[14] This classification has been upheld in authoritative works, including the Flora of North America (Volume 25, 2007), which treats it as the accepted name with no recognized infraspecific taxa in North America.^[6]

Description

Morphology

Distichlis spicata is a perennial, rhizomatous grass that forms dense mats through vegetative propagation. The culms are erect to decumbent, 15 to 90 cm (6 to 35 inches) in height, and are often wiry and rigid. Plants are frequently stoloniferous in addition to rhizomatous, enabling clonal spread and the formation of extensive colonies.^[6]^[4]^[3] The leaves are narrow and stiff, arranged in two vertical ranks along the stem, with blades measuring 1–8 cm long (up to 20 cm occasionally) and 1–5 mm wide. They are flat to folded, ascending to divaricate, and sharp-pointed, often exhibiting salt-crusted tips due to the presence of salt-excreting glands on both surfaces. Leaf sheaths are glabrous, and ligules are short, about 0.5 mm, and erose-ciliate.^[6]^[4]^[15] The root system consists of extensive fibrous roots penetrating up to 70 cm and scaly, monopodial rhizomes that can extend up to 180 cm in length and reach depths of 10-30 cm, though most roots are concentrated in the upper 10–40 cm. This structure facilitates soil stabilization and clonal expansion, with adventitious roots featuring aerenchymatous tissue for gas exchange.^[4]^[3] The inflorescence is a dioecious spike-like panicle, with pistillate panicles congested and 1–8 cm long, and staminate panicles denser and 1-10 cm long. Each raceme bears 2–20 spikelets, which are unisexual and contain 5–20 florets, each 3–6 mm long; lemmas are 3.5–6 mm, and caryopses are awnless, 2–5 mm long.^[6]^[4]^[15]^[16]

Reproduction and growth

Distichlis spicata is a dioecious species, featuring separate male and female plants that reproduce sexually through wind pollination. Male inflorescences produce abundant pollen, which is dispersed by wind to fertilize female flowers, while female inflorescences develop into seeds following successful pollination.^[4] This breeding system contributes to spatial segregation of sexes in populations, influencing reproductive success based on proximity and environmental factors.^[17] Propagation in D. spicata occurs primarily through vegetative means via extensive rhizomes and stolons, enabling rapid clonal spread and colonization of new areas, particularly in challenging saline environments.^[3]^[18] Sexual reproduction supplements this via seeds, though viability is often low in natural settings, with germination rates around 3% in the wild due to dormancy and environmental constraints.^[19] Under saline conditions, seeds exhibit reduced but viable germination, achieving up to 30-50% at moderate salinity levels (150-200 mM NaCl) after treatments like scarification or stratification to break dormancy.^[4]^[20] As a warm-season perennial, D. spicata exhibits peak growth during spring and summer, with active shoot elongation and rhizome extension under favorable temperatures.^[4] Flowering typically occurs from June to October, aligning with warmer months to maximize reproductive output before seasonal dormancy in winter.^[3] The species has a diploid chromosome number of 2n=40, supporting its perennial life cycle and genetic stability across populations.^[21]

Distribution and habitat

Geographic distribution

_Distichlis spicata is native to the Western Hemisphere, with a widespread distribution across North and South America. In North America, it ranges from southern Canada, including provinces such as British Columbia, Alberta, Manitoba, New Brunswick, Nova Scotia, Northwest Territories, Ontario, Prince Edward Island, and Saskatchewan, southward through the United States to Mexico. It is particularly common in the western and southwestern United States, where it occupies diverse saline environments. In South America, the species extends to countries including Ecuador, Peru, Bolivia, Uruguay, Chile, and Argentina.^[6]^[22]^[23] The species has been introduced and naturalized outside its native range, notably in Australia, where it was brought in the 1990s for forage production trials on saline lands in southern regions. Historical spread to such areas likely occurred through human activities, including trade and possibly ship ballast, facilitating its establishment in non-native saline sites. While reports of naturalization in parts of Europe, such as the Mediterranean, and Asia exist, verified distributions remain limited primarily to the Americas and Australia.^[6]^[24] Populations of D. spicata exhibit zonation between coastal and inland areas, with coastal forms often occurring along maritime shorelines and inland variants in arid or semi-arid saline flats and basins. The species tolerates a broad elevational range, from sea level to approximately 2,500 meters (8,200 feet), though extremes up to 2,700 meters have been recorded in some regions. Latitudinally, it spans from about 50°N in southern Canada to roughly 40°S in southern South America, reflecting its adaptability to temperate and subtropical climates.^[4]^[6]

Preferred habitats

Distichlis spicata thrives in saline and alkaline soils, typically with pH values ranging from 6.8 to 9.2, though it can endure extremes up to 10.9. These soils often exhibit salinity levels between 0.03% and 5.4%, with an optimum around 1.5%, corresponding to electrical conductivity (EC) values that support halophytic growth in natural settings. The species favors fine-textured soils with poor drainage, such as those in salt marshes, flats, and meadows, where it tolerates prolonged flooding—over 30 days of submersion—and extended drought periods, with water tables frequently near the surface (16–36 inches deep in some regions).^[4] In terms of climate, Distichlis spicata is suited to temperate and subtropical conditions in USDA hardiness zones 7 through 10, requiring at least 80 frost-free days annually. It performs best in full sun, being shade intolerant, and persists in environments characterized by high evaporation rates and low precipitation, from as little as 0.74 inches in arid basins to higher amounts in coastal areas.^[3]^[4] Distichlis spicata commonly forms monotypic stands in disturbed or hypersaline microhabitats, including coastal dunes, inland playas, and desert scrub, where its rhizomatous growth facilitates dense coverage. It associates with other halophytes in these settings but dominates in zones of elevated salinity and periodic disturbance.^[4]

Ecology

Environmental adaptations

Distichlis spicata exhibits remarkable adaptations to saline environments, primarily through specialized salt-excreting glands located on the leaf surfaces that actively secrete excess ions such as Na⁺, K⁺, and Cl⁻, maintaining internal osmotic balance by extruding salts in a diurnal rhythm with a Na:K ratio of approximately 14:1.^[25] These bicellular glands, consisting of a basal and cap cell, operate under turgor pressure to deposit salts via cuticular cavities, enabling the plant to tolerate soil salinities ranging from 0.03% to 5.4% NaCl.^[25] Additionally, osmotic adjustment occurs through ion compartmentalization, where the plant maintains a near 1:1 Na:K ratio in shoots to regulate water potential and support turgor under high salinity.^[25] Laboratory studies indicate that growth is optimal at around 15,000 ppm soluble salts, corresponding to approximately 1.5% NaCl, beyond which productivity declines due to ion toxicity.^[25]^[4] For drought tolerance, D. spicata relies on its extensive rhizomatous system and fibrous roots that extend up to 28 inches deep, facilitating access to subsurface moisture in arid conditions where annual precipitation may be as low as 0.74 inches.^[4] Inland ecotypes demonstrate high drought resistance, remaining green through extended dry periods, aided by stomatal adaptations such as positioning in leaf furrows and bulliform cells that promote leaf rolling to minimize transpiration.^[25]^[3] In response to fire, the plant resprouts vigorously from rhizomes, establishing rapidly post-burn through lateral spread and seed germination, with cover increasing from 10% to 70% within 43 months in affected salt marshes.^[4] Silica-containing trichomes on leaves contribute to desiccation resistance by reducing water loss and providing structural reinforcement, while also deterring herbivores through abrasiveness.^[25] The species further adapts to other abiotic stresses, including alkalinity, with tolerance to soil pH levels from 6.8 to 10.9, thriving in alkaline flats and sodic sites where it aids in soil rehabilitation.^[4]^[3] Its wiry, low-growing habit and dense rhizome mats confer resistance to trampling, making it suitable for stabilizing heavily trafficked saline areas like cattle trails.^[26]^[3] In nutrient-poor soils, rhizomes penetrate heavy clays and shales, gradually improving soil structure and enabling persistence in low-fertility environments.^[25] As a C4 photosynthetic plant, D. spicata efficiently utilizes high light intensities and elevated temperatures, with optimal germination under a 104°F day/50°F night regime, supporting productivity in hot, arid saline habitats.^[25]^[4]

Biotic interactions

Distichlis spicata engages in both competitive and facilitative interactions with other plant species in marsh ecosystems. It competes with Spartina species for resources such as space and nutrients, often being restricted to lower densities and disturbed habitats by dominant Spartina alterniflora and Spartina patens in coastal salt marshes.^[27] Additionally, as a dioecious species, D. spicata exhibits intersexual competition at the seedling stage, where male and female plants show greater competitive effects on each other than intrasexual interactions, influencing population structure and spatial segregation by sex.^[28] Conversely, D. spicata facilitates other plants, including Spartina seedlings, by stabilizing soil through its extensive rhizome system, reducing erosion and creating suitable microsites for establishment in dynamic marsh environments.^[27] Herbivory impacts D. spicata significantly, with grazing by various wildlife and livestock shaping its distribution and abundance. Geese, such as snow geese, readily consume its rhizomes and roots, particularly after disturbances like fire, while ducks feed on seeds, contributing to seed dispersal.^[4] Livestock, including cattle and sheep, graze it opportunistically on saline soils when preferred forage is scarce, though its coarse foliage results in low overall palatability; big game like deer and pronghorn use it minimally.^[4] In symbiosis, D. spicata forms associations with vesicular-arbuscular mycorrhizal fungi, which enhance nutrient uptake—particularly phosphorus—and maintain favorable potassium-to-sodium ratios in saline soils, supporting growth despite high salt stress.^[29] Within food webs, D. spicata plays a foundational role by providing habitat and resources across trophic levels in salt marsh ecosystems. It offers cover and nesting sites for birds, such as ducks and Wilson's phalaropes, and shelter for invertebrates, fish, and decapod crustacean larvae, bolstering biodiversity in coastal habitats.^[3] Furthermore, as a larval host for skipper butterflies like the obscure skipper (Panoquina panoquinoides), it supports insect populations, while its decomposition contributes organic detritus that fuels detritus-based food chains, indirectly nourishing higher consumers like fish and birds.^[4]^[30]

Human uses

Traditional and cultural uses

Distichlis spicata, commonly known as saltgrass, has been utilized by various indigenous peoples in California for its salt content, derived from specialized glands on the leaves that excrete excess salts. The Kawaiisu people extracted salt by scraping the crystalline deposits from the leaf surfaces to form blocks used as a seasoning and preservative.^[3] Similarly, the Cahuilla burned the leaves to ash to obtain salt for culinary condiments.^[31] Other tribes, such as those in the Kern Valley, harvested the grass during hot, dry periods, spreading it on mats to collect the salt crust for dietary use.^[18] Beyond salt extraction, D. spicata served medicinal purposes among several California indigenous groups. The Kawaiisu prepared decoctions or infusions of the plant to treat dermatological issues like insect bite pimples, rapid heartbeat, constipation as a laxative, and gonorrhea.^[32] The Yokuts used a decoction boiled into gum for colds and to stimulate appetite in cases of loss.^[33] The Cahuilla employed the leaves as a scouring material for cleaning tools or removing cactus thorns, leveraging the plant's abrasive texture.^[34] Additionally, the Kawaiisu made dried grass cakes or green infusions as beverages.^[32] In the 19th century, European-American settlers in the western United States recognized D. spicata for its forage potential, particularly in saline marshes and alkaline plains. It was freely cut alongside other marsh grasses to provide pasturage for livestock, though considered inferior due to its coarse nature and lower nutritional value compared to other species.^[35] Early accounts, such as those from the late 1800s, noted its role in sustaining cattle and horses during dry seasons when it remained green, contributing to rangeland management in arid regions. No widespread ceremonial or symbolic roles have been documented in historical records.

Agricultural and ecological uses

_Distichlis spicata serves as a valuable forage crop in saline environments, where it supports livestock grazing on lands unsuitable for traditional agriculture. Under favorable soil and moisture conditions with saline irrigation, it produces dry matter yields of approximately 9,081 kg/ha and protein yields of 1,300 kg/ha, making it a nutritious option for cattle and sheep during spring, summer, fall, and winter grazing.^[5] This grass is particularly effective for establishing saline pastures, enhancing productivity in salt-affected rangelands without requiring extensive soil amendments.^[3] In ecological management, the extensive rhizome system of D. spicata provides robust erosion control, stabilizing wetland banks and reducing soil loss in saline areas. Its tenacious root network binds erodible soils, slowing overland water flow and preventing degradation in coastal and inland saline habitats.^[36] This attribute has led to its application in reclaiming mined or polluted saline lands, where it improves soil chemical and physical properties while tolerating heavy metal contamination from activities like mining and smelting.^[37] For instance, populations of D. spicata thrive on metal-impacted sites in southwestern Montana, demonstrating tolerance to elevated levels of zinc, copper, and lead.^[37] Beyond forage and stabilization, D. spicata shows promise as a biofuel crop due to its high biomass production in saline conditions, offering a sustainable alternative for marginal lands. Halophyte studies highlight its potential for bioenergy feedstocks, leveraging its adaptation to high-salinity regimes for viable yields without competing with food crops.^[38] Additionally, it aids phytoremediation by accumulating heavy metals in brackish soils, supporting cleanup efforts in contaminated wetlands. In habitat enhancement, D. spicata contributes to salt marsh ecosystems by providing cover and forage for birds, mammals, invertebrates, fish, and marine larvae, thereby bolstering biodiversity in restored saline wetlands.^[36]

Cultivation and conservation

Cultivation methods

Distichlis spicata is primarily propagated vegetatively through rhizome cuttings, which is the more reliable and commonly used method due to the plant's extensive rhizomatous growth habit, e.g., the conservation release 'LK 517f' propagated via rhizomes or plugs on 1-foot centers.^[36] Rhizome cuttings should be collected and planted in early spring, ensuring they remain moist to prevent desiccation; they can be stored for up to 28 days at 35–50°F (2–10°C) and 60–75% relative humidity before planting. Cuttings are planted 1–2 inches (2.5–5 cm) deep and spaced approximately 1 foot (30 cm) apart to allow for clonal spread, with establishment success enhanced by initial irrigation during the first growing season.^[5]^[3] Seed propagation is possible but less straightforward, as natural germination rates are low (around 13%) without pretreatment. Seeds benefit from scarification (mechanical or chemical) and stratification at 39°F (4°C) in moist sand for 30 days, which can increase germination to 54–62% in saline media;^[20]^[39] scarified seeds are sown in coarse, saline soils under full sun conditions for optimal results.^[5] Site preparation for cultivation involves selecting areas with saline or alkaline soils (pH 7.5–8.5) and providing full sun exposure, as the species thrives in brackish marshes, salt flats, or alkaline meadows. Irrigation with brackish or saline water is recommended during establishment to mimic natural conditions, while spacing supports the plant's rhizomatous expansion without competition. Once established, D. spicata requires minimal site amendments due to its tolerance for poor, compacted soils.^[5] Management of cultivated D. spicata is low-maintenance, with the plant exhibiting strong tolerance to grazing and trampling owing to its rhizomatous structure. For forage production, mowing or harvesting can occur from late summer through fall (June–October flowering period), allowing at least 4 inches of regrowth before resuming grazing to prevent overstress; biannual burning in late fall or winter (September–February) may promote vigor in wetter sites. Overwatering should be avoided after establishment to prevent reduced salinity tolerance, though moderate irrigation sustains productivity in arid conditions.^[5]^[3]^[26]

Conservation status

Distichlis spicata is globally secure, with a NatureServe conservation status rank of G5, indicating it is demonstrably secure across its range and not currently at risk of extinction.^[2] The species is assessed as Least Concern on the IUCN Red List, reflecting its widespread distribution and abundance in suitable habitats, though it has not been comprehensively evaluated at the subspecies level. However, certain inland variants, such as those in seasonally inundated potholes and shallow lakes, are locally rare and vulnerable due to habitat loss from altered hydrology and disturbances. The primary threats to wild populations of D. spicata include wetland drainage and hydrological alterations, which reduce available saline habitats, particularly in inland alkaline meadows. Coastal populations face risks from sea-level rise, which can lead to habitat inundation and loss in salt marshes, exacerbating fragmentation for associated species and potentially the grass itself.^[40] Competition from invasive species, such as non-native grasses in converted agricultural lands, further pressures remnant populations, while pollution from nearby development affects water quality in alkaline meadows.^[41] Protection efforts for D. spicata involve its incorporation into restoration projects to bolster wild populations and habitat connectivity, such as in tidal marsh enhancements along coastlines.^[42] The species is monitored within U.S. national parks and wildlife reserves, including areas like Aransas Bay, where habitat restoration supports its persistence amid broader ecosystem threats.^[42] No subspecies are currently listed as endangered, but concerns exist regarding reduced genetic diversity in fragmented habitats.

References

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