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Home , Aquatic plant, Spartina, Spartina densiflora

Smooth Cordgrass ( alterniflora)

Thomas Pham

Fish 423

Aquatic Invasion Ecology

Final Report: Fall 2011

Diagnostic information

Scientific name Order: Family: Genus: Spartina Species: alterniflora

Common names: Smooth cordgrass, saltmarsh cordgrass, saltwater cordgrass, Atlantic cordgrass, oystergrass

Basic identification key

Using the Key to West Spartina Species’ dichotomous key one can properly identify Spartina alterniflora among the other

Spartina species. According to the key S. Figure 1. Spartina alterniflora at a salt in East alterniflora has leaf blades when fresh, Sandwich, Massachusetts. Nelson DeBarros @ internodes that are fleshy, a leaf width that is at USDA-NRCS Database most 25mm, reddish streaks or red pigment often present at the base of young and healthy Life-history and basic ecology shoots. Additionally, the USDA Natural Resources Conservation service describes S. Life cycle, environmental optima and tolerances alterniflora as a grass with long hollow Spartina alterniflora is a perennial rhizomatous . It ranges from 2 to 7 feet tall and has grass that grows in intertidal zones (Subudhi et leaf blades that are around 12-20 inches long. al. 2009). In specific it grows in , mud Lastly, during the months of September and flat, and sand flat . It prefers habitats October, seedheads are present that are around with low or moderate wave action. Furthermore, 12 inches in length and can carry spikes that this is highly adapted to grow in very high carry 12-15 spikelet seeds. concentrations and can grow in up to 60‰ (Bertness 1991). It can grow in a variety of substrates including: sand, silt, cobble, clay, and gravel. S. alterniflora is a spikelets which hold seeds that generally develop in July through October. Pollination is achieved by wind and seeds are dispersed primarily dispersed by water which can carry the seeds long distances due to tides and currents. Vegetative fragmentation is the process in which segments of the plant break off and form a new Figure 2. An inundated Spartina alterniflora marsh plant but is genetically identical to the original. particularly persistent species that tolerate a Lastly, S. alterniflora can also spread clonally varying degree of abiotic conditions. It can by rhizomes. When pieces of root break survive in complete submersion of water for up off, they can sometimes regrow into a new plant to 12 hours and in water with pH levels ranging that is also a genetic clone to the original. from 4.5 to 8.5 (Landin 1991). Nutrient supply is a limiting factor for S. alterniflora. Bursesh et Current geographic distribution al. (1980) found that nitrogen is an important determining factor for growth and Distribution in the United States for S. alterniflora in Louisiana salt . According to the United States Additionally, they observed a greater influence Department of Agriculture Natural Resources of nitrogen added to inland meadows compared Conservation Service Spartina alterniflora to streamside meadow. currently resides among 21 different states

including: Alabama, California, Connecticut, Reproductive strategies Delaware, Florida, Georgia, Louisiana, Spartina alterniflora has three methods Massachusetts, Maryland, Maine, Mississippi, of reproduction that involve both sexual and North Carolina, New Hampshire, New Jersey, clonal processes. It reproduces by seed, rhizome, New York, Oregon, Rhode Island, South or vegetative fragmentation (Daehler and Strong Carolina, Texas, Virginia, and Washington. 1994). It produces inflorescences containing Figure 3. Current distribution of Spartina alterniflora across the United States (USGS)

Spartina alterniflora is non-native to the state of History of invasiveness Washington. There are three main regions of

Washington State that S. alterniflora has Spartina alterniflora is a rhizomatous established populations: Puget Sound, Grays grass native to the Atlantic and Gulf coast Harbor, and Willapa Bay. It is believed that it marshes of (Xiao et al. 2011). It was introduced by accident into Willapa Bay dominates the marshes in its native range. S. around the late nineteenth century as a alterniflora has been introduced to new regions hitchhiker with oysters shipped from the both intentionally and unintentionally. It has Atlantic coast (Dennis et al. 2011). Seeds of S. been introduced to the west coast of the United alterniflora likely were inadvertently brought States, Great Britain, the Atlantic coast of into the same barrels that the oysters were being , and New Zealand (Marchant et al 1970; shipped in. Furthermore, Stiller and Denton Partridge et al 1987; Hitchcock et al. 1969). (1995) performed random amplified Distribution in Washington State polymorphic DNA (RAPD) analysis to determine the genetic history of the S. and wildlife ”. It is currently alterniflora populations in Willapa. Their data classified as a Class A noxious weed meaning suggests that the entire S. alterniflora population that eradication of this plant is required by law. inhabiting the Willapa Bay region descended Unlike the populations in Willapa Bay, S. from a single introduced clonal colony. The alterniflora was intentionally introduced into establishment of this population was spread Puget Sound by landowners. It was introduced primarily by . Rhizome and into Padilla Bay sometime between 1940 and vegetative fragmentation contributed little to the 1946 by the Dike Island Gun Club in order to spread in this region (Civille et al. 2005). S. stabilize the land that the gun club was on alterniflora was recognized as a pest weed in (Parker and Aberle 1979). A hybrid species Willapa Bay and was placed on Washington Spartina anglica is able to hybridize with S. State’s noxious weed list in 1989. The alterniflora and was introduced to Puget Sound Washington State Noxious Weed Control Board in 1961 (Hacker et al. 2001). defines a noxious weed as “the traditional, legal Populations of S. alterniflora also exist in Grays term for any invasive, non-native plant that Harbor. However, the pathway of introduction threatens agricultural crops, local ecosystems of of it is not known for this particular region. In addition to S. alterniflora, is also found here.

Distribution in Oregon S. alterniflora is also present in the State of Oregon, although at much lower densities than observed in Washington State. As of now three infestations of S. alterniflora have been observed in Oregon. One of the colonization’s took place in the Siuslaw in Florence, Oregon. It was intentionally planted around the 1970’s at the Port of Siuslaw (Frenkel 1990, Strong and Ayres 2009). S. alterniflora was also detected in Coos Bay in 2005 at a dredge material disposal site (Oregon Figure 4. Distribution of Spartina alterniflora sites in Response Plan 2007). It is believed that Washington State 2010 (WSDA) unintentional transplantation was the vector for the introduction of it into Coos Bay. Finally, the third and most recent infestation of Factors influencing establishment and spread S. alterniflora in Oregon was discovered in 2008 There are a number of factors that influence the at Youngs Bay (ODA Plant Division Annual establishment of Spartina alterniflora. One Report 2010). However, early detection found factor that actually slowed the rate of invasion of the patch to be only 800 square feet in size and it into the Pacific Northwest was Allee effects was dealt with quickly. As of 2010 no new which is “a positive relationship between any plants have been found in Youngs Bay. component of fitness of a species and density of conspecifics” (Stephens et al. 1999). Davis et al. Invasion process (2004) conducted an experiment and discovered that pollen limitations can cause an Allee effect Pathways, vectors and routes of introduction on S. alterniflora meaning that it can slow its Spartina alterniflora historically has rate of colonization. Additionally, without Allee invaded the Pacific Northwest by both effects S. alterniflora would have spread across intentional and unintentional introductions. The Willapa Bay at a much higher rate (Taylor et al. initial pathway that has led to S. alterniflora’s 2004) and likely would have covered the entire invasion into Washington has been through bay a long time ago (Strong and Ayres 2009). , in particular the stocking of oysters. S. alterniflora was introduced into Willapa Bay Potential ecological and/or economic impacts in the late nineteenth century as part of the Spartina alterniflora is an ecosystem oyster cultivation efforts (Dennis et al. 2011). S. engineer. Jones et al. (1994) coined the term alterniflora seeds hitchhiked along with oyster ecosystem engineer and defined it as: shipments by train from Atlantic marshes where “organisms that directly or indirectly modulate its native region lay. the availability of resources (other than In addition to unintentional introductions, there themselves) to other species, by causing physical have also been intentional introductions of S. state changes in biotic or abiotic materials. In so alterniflora. In its native range, S. alterniflora is doing they modify, maintain and/or create valued for its ability to alleviate erosion habitats”. S. alterniflora alters the ecosystem in (Simenstad and Thom. 1995). The very same which it habitats by a number of different properties have led to intentional introduction means. It can change nutrient cycling, into Washington and Oregon. Furthermore, it hydrology, sediment deposition patterns, and has the ability to trap sediment very well. It has furthermore it can transform open intertidal been introduced into New Zealand because of into elevated meadows filled with this property as a tool for restoration nothing but S. alterniflora (Crooks 2002). These (Partridge 1987). impacts have led to significant change in the landscape of and intertidal zones in the (2000) suggests that a non-native crab Carcinus Pacific Northwest. In the absence of S. maenas are more concentrated in areas where alterniflora estuaries in the Pacific Northwest Spartina are found. are generally gently sloped and shallow, bare S. alterniflora also has the potential to cause mudflats. However, the S. alterniflora can massive economic damage as well. Although it transform them into steep and deep tidal has not occurred, it has the potential to cause channels. Furthermore, the once bare mudflats damage to oyster and commercial . If can become completely covered in meadows of they raise the elevation of the estuaries, they can S. alterniflora. In addition, it has the ability to become unsuitable for oyster aquaculture. increase sedimentation and decrease the effects Oyster farming is a large industry for both of wave action (Gleason et al. 1979) while also Washington and Oregon State and S. alterniflora causing increased flooding. has the potential to cause damage to these S. alterniflora not only affects the abiotic markets by land alterations. Furthermore they structure of communities but can also have can reduce the prey resources for Oncorhynchus impacts on native fauna and flora. It is a robust keta (Chum) in its juvenile stage as well as invader and can out-compete other species such Parophrys vetulus (English sole) both of which as marina (eelgrass). Not only is this are important commercial fish for Washington detrimental to Z. marina but also to the species and Oregon (Noxious Weed Control Board). In that rely on it such various addition S. alterniflora can also can economic including juvenile Dungeness crab damage by altering beaches which are important (Metacarcinus magister) (McMillan et al. 1995). to the tourism markets for Washington and The loss of Z. marina can also have negatively Oregon (Oregon Response Plan 2007). cascading effects on Anas Americana (American wigeon), Anas acuta (Northern Pintail), and Management strategies and control methods Branta bernicla (Brant) all of which rely on Z. marina for forage (Oregon Response Plan 2007). A large amount of time, money, and Furthermore, the large colonization of bare effort has been put into the control and mudflats of S. alterniflora greatly reduces the eradication of Spartina alterniflora in the Pacific open habitat for many different shorebirds and Northwest. In Washington State, management of waterfowl. It is currently listed as a threat to S. alterniflora began in the 1990’s (Hedge et al. birds by the American Bird Conservancy. The 2003). In 1995, the WSDA was put in control of disturbance that S. alterniflora can cause on its its management. In 2003, the Portland State habitat may even open up opportunities for other University Center for Lakes and Reservoirs . A study by and Dumbauld created the Oregon Spartina Response Plan with the goal of “prevent(ing) the establishment and However, Gustafson et al. (2006) tested the spread of any Spartina species in Oregon effects of P. marginata grazing on Spartina estuaries and coastal ”. A number of and found that it is does not exert strong different strategies have been used to control S. top-down control on S. alterniflora. alterniflora. Removal by hand has been used but is limited in a number of ways. It is highly time Literature Cited consuming to remove because care must be taken to remove the entire plant. If residual Bertness MD (1991). Zonation of Spartina rhizomes are left behind, they have the potential Patens and Spartina Alterniflora in New to grow back. This strategy has more England Salt Marsh. Ecology 72: 138-148. effectiveness in controlling younger infestations Buresh RJ, Delaune RD, Patrick WH (1980). than mature (Hedge et al. 2003). Another Nitrogen and Phosphorus Distribution and method that has been used is mowing. Once Utilization by Spartina alterniflora in a again however, this strategy is also limited. It Louisiana Gulf Coast Marsh. Estuaries 3: was found to be neither effective nor cheap as 111-121. sites had to be mowed multiple times to Carr EM, Dumbauld BR (2000). Status of the effectively eliminate it. In addition, European green crab invasion in have also been used. The only authorized Washington coastal estuaries: can for control of S. alterniflora by the expansion be prevented? Journal of Washington Management Shellfish Research 19 : 629-630. Program Environmental Impact Statement is Civille JC, Sayce K, Smith SD, Strong DR Rodeo whose main active ingredient is (2005). Reconstructing a century of glyphosate. There have been large variations in Spartina alterniflora invasion with historical its effectiveness ranging from no effect to records and contemporary remote sensing. complete elimination. The most effective Ecoscience 12: 330-338. method of control in Washington has been a Crooks (2002). Characterizing Ecosystem-Level combination of mowing followed up by Rodeo Consequences of Biological Invasions: The application after it has regrown to 30-45cm, Role of Ecosystem Engineers. OIKOS 97: although it has been found to be highly costly 153-166. (Hedge et al. 2003). One last control method that Daehler CC, Strong DR (1994). Variable has been used is a biological control with the Reproductive Output Among Clones of species Prokelisia marginata (Homopteran plant Spartina alterniflora (Poaceae) Invading hopper). This particular species feeds on San Francisco Bay, California: The Spartina species, specifically its vascular fluids. Influence of Herbivory, Pollination, and Establishment Site. American Journal of Marchant CJ (1970). Evolution in Spartina Botany 81: 307-313. (Gramineae) IV. The cytology of S. Davis HG, Taylor CM, Lambrinos LG, Strong alterniflora Loisel. in North America. DR, Mooney HA (2004). Pollen Botanical Journal of the Linnean Society Limitations Causes an Allee Effect in a 63: 321-326. Wind-Pollinated Invasive Grass (Spartina McMillan RO, Armstrong DA, Dineel PA alterniflora). PNAS: 101: 13804-13807. (1995). Comparison of intertidal habitat Dennis B, Civille JC, Strong DR (2011). Lateral use and growth rates of two northern Puget spread of invasive Spartina alterniflora in Sound cohorts of 0+ age Dungeness crab, uncrowded environments. Biological Cancer magister. Estuaries 18: 390-398. Invasions 13: 401-411. Parker RC, Aberle B (1979). A situation report Gleason ML, Elmer DA, Pien NC, Fisher JS on the Spartina infestation in northwest (1979). Effects of Stem Density upon Washington. Unpublished report to the Sediment Retention by Salt Marsh Cord Washington State Department of game, Grass, Spartina alterniflora Loisel. Mount Vernon Estuaries 2: 271-273. Partridge TR (1987). Spartina in New Zealand. Gustafson DJ, Kilheffer J, Silliman BR (2006). New Zealand Journal of Botany 25: 567- Relative Effects of Littoraria irrorata and 575. Prokelisia marginata on Spartina Simenstad CA, Thom RM (1995). Spartina alterniflora. Estuaries and 29: 639- alterniflora (smooth cordgrass) as an 644. invasive halophyte in Pacific Northwest Hedge P, Kriwoken LK, Patten K (2003). A estuaries. Hortus Northwest 6:9-12, 38-40. Review of Spartina Management in Stiller JW, Denton AL (1995). One hundred Washington State, US. Journal of Aquatic years of Spartina altemiflora (Poaceae) in Plant Management 41: 82-90. Willapa Bay, Washington: random Hitchcock CL, Cronquist A, Ownbey M (1969) amplified polymorphic DNA analysis of an Vascular Plants of the Pacific Northwest. invasive population. Molecular Ecology 4: Part 1: Vascular Cryptogams, 355-363. Gymnosperms, and . Stephens PA, Sutherland WJ, Freckleton RP University of Washington Press, Seattle, (1999). What is the Allee effect? OIKOs Washington 87: 185-190. Jones CG, Lawton JH, Shachak M (1994). Strong DR, Ayres DA (2009) Spartina Organisms as ecosystem engineers. OIKOS Introductions and Consequences in Salt 69: 373-386. Marshes. In: Silliman BR (ed), Grosholz ED (ed), Bertness MD (ed) Human Impacts http://agr.wa.gov/PlantsInsects/Weeds/Spartina/ on Salt Marshes: A Global Perspective, 1st docs/SpartinaReport2010.pdf edn. University of California press, Berkeley, CA. Oregon Department of Agriculture Plant Subudhi PK, Baisakh N (2011). Spartina Division Annual Report 2010 alterniflora Loisel., a halophyte grass model http://www.oregon.gov/ODA/PLANT/docs/pdf/ to dissect salt stress tolerance. In Vitro plant_annual_report_2010.pdf?ga=t Cellular & Developmental Biology – Plant 47: 441-457. West Coast Governors’ Agreement on Taylor CM, Davis HG, Civille JC, Grevstad FS, Health Spartina Eradication Action Hastings A (2004). Consequences of an Coordination Team Work Plan Allee effect in the invasion of a pacific http://westcoastoceans.gov/Docs/Spartina_Final estuary by Spartina alterniflora. Ecology _Work_Plan.pdf 85: 3254-3266. Xiao Y, Tang J, Qing H, Zhou C, An S (2011). Expert contact information in PNW Effects of salinity and clonal integration on growth and sexual reproduction of the Kathleen Sayce invasive grass Spartina alterniflora. Flora P.O. Box 91 206: 736-741. Nahcotta, WA 98637 Phone: (360) 665-5292 (H), (360) 642-1166 (W) Other key sources of information and [email protected] bibliographies Vanessa Howard Written Findings of the Washington State Center for Lake and Reservoirs Noxious Weed Control Board (1995) Portland State University http://www.nwcb.wa.gov/siteFiles/Spartina_alter P.O. Box 751 niflora.pdf Portland, OR 97207-0751 Phone: (503) 725-9076 Oregon Spartina Response Plan 2007 Fax: (503) 725-3834 http://www.clr.pdx.edu/docs/OSRP.pdf [email protected]

Spartina Eradication Program 2010 Progress Nancy Ness Report Grays Harbor County Noxious Weed Board Coordinator P.O. Box R Spartina alterniflora. Since 1995, the Elma, WA 98541 Washington State Department of Agriculture has Phone: (360) 482-2265 spearheaded the eradication of Spartina species. Fax: (360) 482-2662 They have coordinated a number of stakeholders [email protected] and entities in working together to manage Spartina. They have seen great success in Kyle Murphy eliminating S. alterniflora as well as other WSDA Spartina Coordinator Spartina species in the state. From a record high P.O. Box 42560 of 9,260 acres of Spartina spp. observed in Olympia, WA 98504 2003, the WSDA has reduced that amount to 27 Phone: (360) 902-1923 acres as of 2010 (Spartina Eradication Program [email protected] 2010 Progress Report). The effort continues although they believe that the last few acres will Current research and management efforts be the most difficult to remove. The WSDA estimates that 7 solid acres of Spartina will The state of Washington has devoted a remain in Willapa Bay in 2011, less than 0.05 large amount of time and resources to eradicate acres in Grays Harbor, and 5 acres in Puget

Figure 5. Solid acres of Spartina by year statewide based on WSDA estimates. The blue line represents historic Spartina infestations since 2003. The red line indicates the projected Spartina infestation level through 2014. Projection assumes continued funding. Sound. Currently, they are continuing to use integrated pest management techniques such as mechanical, chemical, manual control, or a combination of them as previously described, although more effort is being put into shoreline surveillance as the numbers of acres of Spartina have been greatly reduced. On September 18, 2006 the governors of California, Oregon, and Washington announced the West Coast Governors’ Agreement on Ocean Health. Through this agreement they called for collaboration to manage and protect the ocean and coastal resources along the West Coast. Through this agreement a Spartina eradication program was developed with the goal of eliminating non-native Spartina off of the West Coast by 2018. This program created a comprehensive work plan and outlined a number of tasks in order to achieve the goal of complete eradication. According to the work plan, they are working on developing an internet based GIS (geographic information system) to define the areas in which Spartina has been eradication. Overall this agreement has combined the efforts of California, Oregon, and Washington in eliminating Spartina spp. With the bulk of the infestations managed, the remaining work is synthesizing the collective efforts between agencies and states.