<str<strong>on</strong>g>Proceedings</str<strong>on</strong>g> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>Third</str<strong>on</strong>g> <str<strong>on</strong>g>Internati<strong>on</strong>al</str<strong>on</strong>g> <str<strong>on</strong>g>C<strong>on</strong>ference</str<strong>on</strong>g> <strong>on</strong> <strong>Invasive</strong> SpartinaChapter 1: Spartina BiologyCALIFORNIA CORDGRASS (SPARTINA FOLIOSA), AN ENDEMIC OF SALT MARSH HABITATSALONG THE PACIFIC COAST OF WESTERN NORTH AMERICAM.C. VASEYDepartment <str<strong>on</strong>g>of</str<strong>on</strong>g> Biology, San Francisco State University, San Francisco, CA 94132; andDepartment <str<strong>on</strong>g>of</str<strong>on</strong>g> Envir<strong>on</strong>mental Studies, University <str<strong>on</strong>g>of</str<strong>on</strong>g> California, Santa Cruz, CA 95064; mvasey@sfsu.eduCalifornia cordgrass (Spartina foliosa) is an endemic <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> California Floristic Province in westernNorth America. This paper reviews its historic and c<strong>on</strong>temporary distributi<strong>on</strong>, pr<str<strong>on</strong>g>of</str<strong>on</strong>g>iles its habitatcharacteristics including key adaptive traits, and makes <str<strong>on</strong>g>the</str<strong>on</strong>g> case that S. foliosa should be recognizedas a foundati<strong>on</strong> species within salt marshes characteristic <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> California Floristic Province. Thisinformati<strong>on</strong> is <str<strong>on</strong>g>the</str<strong>on</strong>g>n c<strong>on</strong>sidered in <str<strong>on</strong>g>the</str<strong>on</strong>g> c<strong>on</strong>text <str<strong>on</strong>g>of</str<strong>on</strong>g> introgressive hybridizati<strong>on</strong> between S. foliosa and<str<strong>on</strong>g>the</str<strong>on</strong>g> closely related Spartina alterniflora. Clearly, <str<strong>on</strong>g>the</str<strong>on</strong>g> hybrid between S. foliosa and S. alterniflora isprogressively spreading genes <str<strong>on</strong>g>of</str<strong>on</strong>g> S. alterniflora into pure stands <str<strong>on</strong>g>of</str<strong>on</strong>g> S. foliosa throughout <str<strong>on</strong>g>the</str<strong>on</strong>g> SanFrancisco Bay estuary. While this raises a c<strong>on</strong>cern about <str<strong>on</strong>g>the</str<strong>on</strong>g> potential “extincti<strong>on</strong>” <str<strong>on</strong>g>of</str<strong>on</strong>g> S. foliosa as adistinct genotype, it is suggested that a greater problem may be <str<strong>on</strong>g>the</str<strong>on</strong>g> ecological implicati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> thisintrogressi<strong>on</strong> for salt marshes in <str<strong>on</strong>g>the</str<strong>on</strong>g> San Francisco Bay estuary. Ra<str<strong>on</strong>g>the</str<strong>on</strong>g>r than eradicati<strong>on</strong> per se, it issuggested that focused c<strong>on</strong>tainment may be <str<strong>on</strong>g>the</str<strong>on</strong>g> best strategy for minimizing <str<strong>on</strong>g>the</str<strong>on</strong>g> impact <str<strong>on</strong>g>of</str<strong>on</strong>g> thishybrid <strong>on</strong> salt marshes in <str<strong>on</strong>g>the</str<strong>on</strong>g> California Floristic Province <str<strong>on</strong>g>of</str<strong>on</strong>g> this regi<strong>on</strong>. Fur<str<strong>on</strong>g>the</str<strong>on</strong>g>r, c<strong>on</strong>tinued adaptivemanagement studies that evaluate <str<strong>on</strong>g>the</str<strong>on</strong>g> impacts <str<strong>on</strong>g>of</str<strong>on</strong>g> this species <strong>on</strong> salt marsh restorati<strong>on</strong> arerecommended.INTRODUCTIONIn this review, I will examine what we know aboutCalifornia cordgrass and attempt to frame this discussi<strong>on</strong> in<str<strong>on</strong>g>the</str<strong>on</strong>g> c<strong>on</strong>text <str<strong>on</strong>g>of</str<strong>on</strong>g> c<strong>on</strong>servati<strong>on</strong> management challenges thatinvasive Spartina alterniflora pose to <str<strong>on</strong>g>the</str<strong>on</strong>g> San Francisco Bayestuary. The approach will be to discuss what is known <str<strong>on</strong>g>of</str<strong>on</strong>g><str<strong>on</strong>g>the</str<strong>on</strong>g> historic and c<strong>on</strong>temporary distributi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartinafoliosa, to focus <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> suite <str<strong>on</strong>g>of</str<strong>on</strong>g> adaptive traits that makes S.foliosa such an important comp<strong>on</strong>ent <str<strong>on</strong>g>of</str<strong>on</strong>g> low marsh habitatsin west coast estuaries, and to pr<str<strong>on</strong>g>of</str<strong>on</strong>g>ile S. foliosa as a“foundati<strong>on</strong> species” in this envir<strong>on</strong>ment. I will <str<strong>on</strong>g>the</str<strong>on</strong>g>nsummarize this informati<strong>on</strong> in <str<strong>on</strong>g>the</str<strong>on</strong>g> c<strong>on</strong>text <str<strong>on</strong>g>of</str<strong>on</strong>g> threats posedby hybridizati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> S. foloiosa with <str<strong>on</strong>g>the</str<strong>on</strong>g> invasive Spartinaalterniflora.HISTORIC AND CONTEMPORARY DISTRIBUTION OFSPARTINA FOLIOSAMacD<strong>on</strong>ald and Barbour (1974) c<strong>on</strong>ducted a survey <str<strong>on</strong>g>of</str<strong>on</strong>g>salt marsh vegetati<strong>on</strong> al<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g> North American Pacificcoast ranging from Point Barrow, Alaska to Cabo San Lucas,Baja California. In general, <str<strong>on</strong>g>the</str<strong>on</strong>g>y found three dominant saltmarsh vegetati<strong>on</strong> communities over this extensive range: anarctic, boreal, and north temperate assemblage is dominatedby nor<str<strong>on</strong>g>the</str<strong>on</strong>g>rn grasses such as Deschampsia caespitosa andsedge species such as Carex lygnbyei, which ranges from <str<strong>on</strong>g>the</str<strong>on</strong>g>Seward Peninsula down to Drake’s Estero in Marin County;a south temperate and subtropical assemblage ranges fromSan Francisco Bay to Laguna San Ignacio in Baja California,which is <str<strong>on</strong>g>the</str<strong>on</strong>g> assemblage occupied by S. foliosa; and south <str<strong>on</strong>g>of</str<strong>on</strong>g>this regi<strong>on</strong>, tropical mangrove forests and scrub assemblagesdominate estuarine tidal wetlands.The historic distributi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> S. foliosa occurs in thisintermediate z<strong>on</strong>e, also known as <str<strong>on</strong>g>the</str<strong>on</strong>g> California FloristicProvince (Raven and Axelrod 1978). The California FloristicProvince occurs al<strong>on</strong>g a cism<strong>on</strong>tane regi<strong>on</strong> from sou<str<strong>on</strong>g>the</str<strong>on</strong>g>rnOreg<strong>on</strong> down through nor<str<strong>on</strong>g>the</str<strong>on</strong>g>rn Baja California. It ischaracterized by a Mediterranean-type climate c<strong>on</strong>sisting <str<strong>on</strong>g>of</str<strong>on</strong>g>l<strong>on</strong>g, hot and dry summers punctuated by short, wet and coldwinters. The California Floristic Province has recently beenrecognized as <strong>on</strong>e <str<strong>on</strong>g>of</str<strong>on</strong>g> twenty-five global “biodiversity hotspots,” with over 48% <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> plant species being endemics(Myers et al. 2000; Calsbeek et al. 2003). Spartina foliosa isam<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g>se endemic species.The particular distributi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> S. foliosa occurs in twomajor disjunct regi<strong>on</strong>s: (1) nor<str<strong>on</strong>g>the</str<strong>on</strong>g>rn California (SanFrancisco Bay regi<strong>on</strong>) and, approximately 565 kilometersaway, (2) a series <str<strong>on</strong>g>of</str<strong>on</strong>g> geographically proximal populati<strong>on</strong>sfrom Orange County in sou<str<strong>on</strong>g>the</str<strong>on</strong>g>rn California through southcentralBaja California. The historic nor<str<strong>on</strong>g>the</str<strong>on</strong>g>rn Californiadistributi<strong>on</strong> has been c<strong>on</strong>fused by <strong>on</strong>e case <str<strong>on</strong>g>of</str<strong>on</strong>g>misidentificati<strong>on</strong> and two o<str<strong>on</strong>g>the</str<strong>on</strong>g>r cases <str<strong>on</strong>g>of</str<strong>on</strong>g> relatively recentnor<str<strong>on</strong>g>the</str<strong>on</strong>g>rn range extensi<strong>on</strong>s. The case <str<strong>on</strong>g>of</str<strong>on</strong>g> misidentificati<strong>on</strong> wasfirst recognized by P. Faber (pers<strong>on</strong>al communicati<strong>on</strong>) whoperceived that <str<strong>on</strong>g>the</str<strong>on</strong>g> extensive stands <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina l<strong>on</strong>grecognized in Humboldt Bay wetlands were actuallySpartina densiflora (a caespitose species from SouthAmerica) ra<str<strong>on</strong>g>the</str<strong>on</strong>g>r than S. foliosa. Later, Spicher and Josselyn(1985) published a c<strong>on</strong>firmati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> this observati<strong>on</strong>.Barnhart et al. (1992) speculate that S. densiflora may havebeen introduced to Humboldt Bay as l<strong>on</strong>g ago as <str<strong>on</strong>g>the</str<strong>on</strong>g> 1860sas part <str<strong>on</strong>g>of</str<strong>on</strong>g> shingle or dry ballast deposited during <str<strong>on</strong>g>the</str<strong>on</strong>g> height<str<strong>on</strong>g>of</str<strong>on</strong>g> shipping activities in this formerly bustling port. Spicher-3-
Chapter 1: Spartina Biology<str<strong>on</strong>g>Proceedings</str<strong>on</strong>g> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>Third</str<strong>on</strong>g> <str<strong>on</strong>g>Internati<strong>on</strong>al</str<strong>on</strong>g> <str<strong>on</strong>g>C<strong>on</strong>ference</str<strong>on</strong>g> <strong>on</strong> <strong>Invasive</strong> Spartinaand Josselyn (1985) also reported a local populati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> S.foliosa at Bodega Bay; however, this populati<strong>on</strong> was almostcertainly not present when Barbour (1970) and MacD<strong>on</strong>aldand Barbour (1974) reported <strong>on</strong> salt marsh vegetati<strong>on</strong> inBodega Bay wetlands. In <str<strong>on</strong>g>the</str<strong>on</strong>g> 1990s, a large populati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> S.foliosa was observed col<strong>on</strong>izing <str<strong>on</strong>g>the</str<strong>on</strong>g> accreting delta <str<strong>on</strong>g>of</str<strong>on</strong>g>Lagunitas Creek in sou<str<strong>on</strong>g>the</str<strong>on</strong>g>rn Tomales Bay (Baye 2004,pers<strong>on</strong>al communicati<strong>on</strong>). Howell (1949, 1970) in his flora<str<strong>on</strong>g>of</str<strong>on</strong>g> Marin, however, does not menti<strong>on</strong> this locality for S.foliosa nor do any historic herbarium collecti<strong>on</strong>s suggest thatei<str<strong>on</strong>g>the</str<strong>on</strong>g>r <str<strong>on</strong>g>the</str<strong>on</strong>g> Bodega Bay or Tomales Bay populati<strong>on</strong>s occurredprior to <str<strong>on</strong>g>the</str<strong>on</strong>g> latter part <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> twentieth century. MacD<strong>on</strong>aldand Barbour (1974) specifically remark up<strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> absence <str<strong>on</strong>g>of</str<strong>on</strong>g>S. foliosa in Tomales Bay.On <str<strong>on</strong>g>the</str<strong>on</strong>g> o<str<strong>on</strong>g>the</str<strong>on</strong>g>r hand, Howell (1949, 1970) and historicherbarium collecti<strong>on</strong>s do c<strong>on</strong>firm that S. foliosa hashistorically been found in Drake’s Estero in sou<str<strong>on</strong>g>the</str<strong>on</strong>g>rn PointReyes as well as <str<strong>on</strong>g>the</str<strong>on</strong>g> relatively nearby Bolinas Lago<strong>on</strong> inMarin County. O<str<strong>on</strong>g>the</str<strong>on</strong>g>r than <str<strong>on</strong>g>the</str<strong>on</strong>g>se two outer coastal estuaries,all o<str<strong>on</strong>g>the</str<strong>on</strong>g>r historic nor<str<strong>on</strong>g>the</str<strong>on</strong>g>rn California populati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> S. foliosaare found within <str<strong>on</strong>g>the</str<strong>on</strong>g> lower porti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> San Francisco Bayestuary, ranging from tidal wetlands <str<strong>on</strong>g>of</str<strong>on</strong>g> South San FranciscoBay up through San Pablo Bay and <str<strong>on</strong>g>the</str<strong>on</strong>g> lower tidal wetlands<str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Petaluma River, S<strong>on</strong>oma Creek, and <str<strong>on</strong>g>the</str<strong>on</strong>g> Napa River.Spartina foliosa c<strong>on</strong>tinues up into <str<strong>on</strong>g>the</str<strong>on</strong>g> Carquinez Straightsbut is largely unknown from Suisun Bay. In this easternregi<strong>on</strong>, it is apparently c<strong>on</strong>strained by low brackish marshvegetati<strong>on</strong> dominated by Schoenoplectus acutus andSchoenoplectus californicus and its distributi<strong>on</strong> is dynamicdepending <strong>on</strong> decadal shifts in <str<strong>on</strong>g>the</str<strong>on</strong>g> salinity gradient in thisporti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> estuary. It is somewhat remarkable that <str<strong>on</strong>g>the</str<strong>on</strong>g>reare no historic or current occurrences <str<strong>on</strong>g>of</str<strong>on</strong>g> S. foliosa from <str<strong>on</strong>g>the</str<strong>on</strong>g>Golden Gate south al<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g> central California coast to PointC<strong>on</strong>cepti<strong>on</strong>, including likely tidal wetlands such as ElkhornSlough and Morro Bay. Peter Baye (pers<strong>on</strong>alcommunicati<strong>on</strong> 2004) suggests that before <str<strong>on</strong>g>the</str<strong>on</strong>g> c<strong>on</strong>structi<strong>on</strong><str<strong>on</strong>g>of</str<strong>on</strong>g> jetties, <str<strong>on</strong>g>the</str<strong>on</strong>g>se central Californian lago<strong>on</strong>s were not tidalduring summers (due to low flows) so it is possible that <str<strong>on</strong>g>the</str<strong>on</strong>g>ydid not provide suitable habitat.In summary, S. foliosa in nor<str<strong>on</strong>g>the</str<strong>on</strong>g>rn California washistorically c<strong>on</strong>centrated within <str<strong>on</strong>g>the</str<strong>on</strong>g> lower, saline reaches <str<strong>on</strong>g>of</str<strong>on</strong>g><str<strong>on</strong>g>the</str<strong>on</strong>g> San Francisco Bay estuary where it was an importantc<strong>on</strong>stituent <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> extensive salt marshes that historicallydominated tidal wetlands in this regi<strong>on</strong>. It appears to beslowly moving north al<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g> coast and is now present inBodega Bay. Its eastern distributi<strong>on</strong> in <str<strong>on</strong>g>the</str<strong>on</strong>g> San FranciscoBay estuary is dynamic and dependant <strong>on</strong> l<strong>on</strong>g-term shifts <str<strong>on</strong>g>of</str<strong>on</strong>g><str<strong>on</strong>g>the</str<strong>on</strong>g> salinity gradient in <str<strong>on</strong>g>the</str<strong>on</strong>g> Carquinez Straight and lowerSuisun Bay subregi<strong>on</strong>.The sec<strong>on</strong>d historic center <str<strong>on</strong>g>of</str<strong>on</strong>g> distributi<strong>on</strong> for S. foliosais relatively well documented by MacD<strong>on</strong>ald and Barbour(1974), Trnka and Zedler (2000), and Wiggins (1980). Thenor<str<strong>on</strong>g>the</str<strong>on</strong>g>rnmost historic locality is Mugu Lago<strong>on</strong> (OrangeCounty), c<strong>on</strong>siderably south <str<strong>on</strong>g>of</str<strong>on</strong>g> Point C<strong>on</strong>cepti<strong>on</strong>. There is<str<strong>on</strong>g>the</str<strong>on</strong>g>n a fairly c<strong>on</strong>tinuous occurrence <str<strong>on</strong>g>of</str<strong>on</strong>g> S. foliosa whereverc<strong>on</strong>diti<strong>on</strong>s are suitable down to <str<strong>on</strong>g>the</str<strong>on</strong>g> Tijuana Estuary in SanDiego County (e.g. Anaheim Bay, Bolsa Chica Bay,Newport Bay, Santa Margarita River, Los PeñasquitosLago<strong>on</strong>, San Elijo Lago<strong>on</strong>, San Diegito Lago<strong>on</strong>, Missi<strong>on</strong>Bay, and San Diego Bay). This distributi<strong>on</strong> c<strong>on</strong>tinues in arelatively c<strong>on</strong>sistent fashi<strong>on</strong> al<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g> coast <str<strong>on</strong>g>of</str<strong>on</strong>g> BajaCalifornia in Estero de Punta Banda, Bahia de San Quintin,Laguna Guerrero Negro, Ojo de Liebre, Laguna San Ignacio,and Bahia de la Magdalena. It is <str<strong>on</strong>g>of</str<strong>on</strong>g> interest that Bahia de laMagdalena is at a boundary between tropical and subtropicalmarine ecosystems. It may be that S. foliosa is able to extendfur<str<strong>on</strong>g>the</str<strong>on</strong>g>r south than most California Floristic Province plantendemics because it is resp<strong>on</strong>ding as much to <str<strong>on</strong>g>the</str<strong>on</strong>g>se marineinfluences as to Mediterranean climate influences. South <str<strong>on</strong>g>of</str<strong>on</strong>g>Laguna Guerrero Negro, S. foliosa begins to co-occur withmangroves and mangrove-like shrubs such as Rhizophoramangle, Laguncularia racemosa, C<strong>on</strong>ocarpus erecta, andAvicennia germinans. MacD<strong>on</strong>ald and Barbour (1974) pointout that <str<strong>on</strong>g>the</str<strong>on</strong>g>se same mangrove associates co-occur withsmooth cordgrass (S. alterniflora) in marshes near Tabasco<strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> Gulf <str<strong>on</strong>g>of</str<strong>on</strong>g> Mexico. Thus, <str<strong>on</strong>g>the</str<strong>on</strong>g> closest geographic distancebetween California cordgrass and smooth cordgrass is at S.foliosa’s sou<str<strong>on</strong>g>the</str<strong>on</strong>g>rn distributi<strong>on</strong>al limit. There are <strong>on</strong>ly about1,200 kilometers separating <str<strong>on</strong>g>the</str<strong>on</strong>g> Gulf coast <str<strong>on</strong>g>of</str<strong>on</strong>g> Mexico from<str<strong>on</strong>g>the</str<strong>on</strong>g> Pacific coast <str<strong>on</strong>g>of</str<strong>on</strong>g> Baja California.HABITAT CHARACTERISTICS AND ADAPTIVE TRAITSThroughout its entire range, S. foliosa occupies adistinctive z<strong>on</strong>e associated with coastal salt marshes. In mostcases, it is <str<strong>on</strong>g>the</str<strong>on</strong>g> <strong>on</strong>ly vascular plant species present in thisz<strong>on</strong>e, which roughly corresp<strong>on</strong>ds to bay or channel marginsat or near <str<strong>on</strong>g>the</str<strong>on</strong>g> mean high tide line (Schoenherr 1995). Everyday, S. foliosa habitat is inundated by salt water for <strong>on</strong>e toseveral hours. Fur<str<strong>on</strong>g>the</str<strong>on</strong>g>r, it is exposed to <str<strong>on</strong>g>the</str<strong>on</strong>g> forces <str<strong>on</strong>g>of</str<strong>on</strong>g> waveacti<strong>on</strong> and high velocity channel flows. This combinati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g>high salinity, prol<strong>on</strong>ged inundati<strong>on</strong>, and daily hydrologicaldisturbance is obviously bey<strong>on</strong>d <str<strong>on</strong>g>the</str<strong>on</strong>g> tolerance limits foro<str<strong>on</strong>g>the</str<strong>on</strong>g>r salt marsh vascular plant species. Spartina foliosa isable to tolerate <str<strong>on</strong>g>the</str<strong>on</strong>g>se extreme c<strong>on</strong>diti<strong>on</strong>s because <str<strong>on</strong>g>of</str<strong>on</strong>g> a uniquesuite <str<strong>on</strong>g>of</str<strong>on</strong>g> adaptive traits that include: (1) C4 photosyn<str<strong>on</strong>g>the</str<strong>on</strong>g>sis, aphysiological drought-adaptive mechanism which meansthat about half as much water is needed for <str<strong>on</strong>g>the</str<strong>on</strong>g> same amount<str<strong>on</strong>g>of</str<strong>on</strong>g> carbohydrate produced; (2) epidermal salt glands whichalso help to c<strong>on</strong>serve water by reducing salt c<strong>on</strong>centrati<strong>on</strong>sin cell sap; (3) possessi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> deep rhizomes (about 15 cm)which regenerate new segments each seas<strong>on</strong> and createextensive, branching mats that penetrate anaerobic sedimentsand anchor shoreline habitat; (4) stems and leaves that arecomposed <str<strong>on</strong>g>of</str<strong>on</strong>g> aerenchymous tissue that allows oxygen totravel from aerial porti<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> plant down to <str<strong>on</strong>g>the</str<strong>on</strong>g> rhizomesthat are embedded in anaerobic sediments; and (5) stems andleaves that are about <strong>on</strong>e meter tall, enabling vegetativestructures to remain above extreme high tide levels so thatoxygen can reliably be transported to <str<strong>on</strong>g>the</str<strong>on</strong>g> roots.-4-
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