<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 BiologyEVOLVING INVASIBILITY OF EXOTIC SPARTINA HYBRIDS IN UPPER SALT MARSH ZONESOF SAN FRANCISCO BAYM.R. Pakenham-Walsh 1 , D.R. Ayres 2 , and D.R. Str<strong>on</strong>g 21 Regulatory Divisi<strong>on</strong>, U.S. Army Corps <str<strong>on</strong>g>of</str<strong>on</strong>g> Engineers, 1325 J Street, Room 1480, Sacramento, CA 958142 Secti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> Evoluti<strong>on</strong> and Ecology, University <str<strong>on</strong>g>of</str<strong>on</strong>g> California, Davis, One Shields Avenue, Davis, CA 95616Keywords: Spartina, Sarcocornia virginica, invasive, hybridizati<strong>on</strong>INTRODUCTIONInvasi<strong>on</strong> by hybrid cordgrass (Spartina alterniflora xSpartina foliosa) is pr<str<strong>on</strong>g>of</str<strong>on</strong>g>oundly altering habitat structurewithin <str<strong>on</strong>g>the</str<strong>on</strong>g> intertidal z<strong>on</strong>e <str<strong>on</strong>g>of</str<strong>on</strong>g> San Francisco Bay, California.Spartina hybrids exhibit wide ecological tolerance comparedto native S. foliosa and are invading <str<strong>on</strong>g>the</str<strong>on</strong>g> naturallyunvegetated lower intertidal z<strong>on</strong>e. Heterogeneous hybridgenotypes exhibit traits <str<strong>on</strong>g>of</str<strong>on</strong>g> both salinity tolerance andcompetitive vigor, which may enable invasi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> highermarsh z<strong>on</strong>es historically dominated by <str<strong>on</strong>g>the</str<strong>on</strong>g> highly salttolerantnative pickleweed species Sarcocornia virginica.The hybrid cordgrass swarm possesses a high degree <str<strong>on</strong>g>of</str<strong>on</strong>g>genetic variati<strong>on</strong>, with bi-directi<strong>on</strong>al introgressi<strong>on</strong> due tooverlapping flowering periods <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrids and both parentalspecies (Ayres et al. 1999; Antilla et al. 2000). Hybridizati<strong>on</strong>has been shown to play a role in large and rapid adaptiveevoluti<strong>on</strong> (Rieseberg et al. 2003), promoting <str<strong>on</strong>g>the</str<strong>on</strong>g> ability forniche separati<strong>on</strong> between hybrids and parental species(Rieseberg et al. 1999).Drawing <strong>on</strong> field observati<strong>on</strong>s indicating higher relativefitness <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrid Spartina compared to native S. foliosa in<str<strong>on</strong>g>the</str<strong>on</strong>g> higher marsh z<strong>on</strong>es, we c<strong>on</strong>ducted two experiments toaddress <str<strong>on</strong>g>the</str<strong>on</strong>g> threat <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrid col<strong>on</strong>izati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> Sa. virginicahabitat. A greenhouse experiment investigated salinitytolerance <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrids. A field experiment examinedcompetitive suppressi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrids by Sa. virginica.METHODSFor <str<strong>on</strong>g>the</str<strong>on</strong>g> greenhouse experiment, cl<strong>on</strong>al fragments <str<strong>on</strong>g>of</str<strong>on</strong>g>eighteen hybrid genotypes were collected from CogswellMarsh, Hayward, California. In additi<strong>on</strong> to <str<strong>on</strong>g>the</str<strong>on</strong>g> hybrids, <strong>on</strong>eS. alterniflora and two S. foliosa genotypes (21 plants total)were grown at three salinity levels (10, 25 and 40 ppt) withthree replicati<strong>on</strong>s for <strong>on</strong>e growing seas<strong>on</strong> in Davis,California (Fig. 1). Total dry biomass (roots and abovegroundmaterial) was determined at <str<strong>on</strong>g>the</str<strong>on</strong>g> end <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> seas<strong>on</strong>,and effects <str<strong>on</strong>g>of</str<strong>on</strong>g> genotype, salinity and <str<strong>on</strong>g>the</str<strong>on</strong>g>ir potentialinteracti<strong>on</strong> determined with statistical analysis.The field experiment was c<strong>on</strong>ducted at Cogswell Marsh,a former salt p<strong>on</strong>d opened to tidal acti<strong>on</strong> in 1980. Four plottreatments (pure Spartina sp., pure Sa. virginica, clipped Sa.virginica and unclipped Sa. virginica; five replicates) wereset up at each <str<strong>on</strong>g>of</str<strong>on</strong>g> eight hybrid cl<strong>on</strong>e locati<strong>on</strong>s and <strong>on</strong>e S.foliosa cl<strong>on</strong>e (Fig. 2). End-<str<strong>on</strong>g>of</str<strong>on</strong>g>-seas<strong>on</strong> shoot density andabove-ground biomass were determined for each plot, andeffects <str<strong>on</strong>g>of</str<strong>on</strong>g> genotype and plot type were determined withstatistical analysis.Fig. 1., left.40 ppt. (left) vs.10 ppt. (right)in salinityexperiment.Fig. 2., right.Locati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g>nine Spartinagenotypes atCogswell Marsh.-29-
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> Spartina180.00160.00140.00Total Dry Biomass (g)120.00100.0080.0060.0040.0020.000.002-15(foli)S. foli 1-9 S. alt 3-7 3-2 3-4 2-5 2-17 1-5 2-13 3-13 3-1 3-10 3-5 2-6 1-17 2-8 2-16 3-8 2-11Spartina GenotypeLow Medium HighFig. 3. Total biomass <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina genotypes at each salinity level, in increasing order <str<strong>on</strong>g>of</str<strong>on</strong>g> total biomass achieved by each genotype(low + medium + high salinities).RESULTSHybrid Spartina genotypes exhibited great variability inmorphological traits, resp<strong>on</strong>se to salinity stress andcompetitive suppressi<strong>on</strong> by Sa. virginica. Salinity reducedtotal biomass <str<strong>on</strong>g>of</str<strong>on</strong>g> most Spartina genotypes. Several hybridcordgrass genotypes exhibited more robust growth than <str<strong>on</strong>g>the</str<strong>on</strong>g>parental species at both low and high salinity levels (Fig. 3).Results <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> field experiment indicated that shootdensity <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrid cordgrasses was highly sensitive to Sa.virginica removal, increasing an average <str<strong>on</strong>g>of</str<strong>on</strong>g> 105% (range =39-211%; Table 1). End-<str<strong>on</strong>g>of</str<strong>on</strong>g>-seas<strong>on</strong> biomass <str<strong>on</strong>g>of</str<strong>on</strong>g> cordgrassgenotypes increased an average <str<strong>on</strong>g>of</str<strong>on</strong>g> 64% in resp<strong>on</strong>se to Sa.virginica removal (range = 2-144%; Table 1). Genotypes inTable 1 are listed in order <str<strong>on</strong>g>of</str<strong>on</strong>g> appearance <strong>on</strong> Fig. 3.CONCLUSIONSCertain hybrids (e.g., genotypes 1-5 and 2-8) showed acombinati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> competitive vigor in <str<strong>on</strong>g>the</str<strong>on</strong>g> field and relativelystr<strong>on</strong>ger performance in higher salinity c<strong>on</strong>diti<strong>on</strong>s,exhibiting <str<strong>on</strong>g>the</str<strong>on</strong>g> potential to successfully expand into highermarsh z<strong>on</strong>es. The native S. foliosa has weaker competitiveabilities and tolerance for salinity. Our results support apositive associati<strong>on</strong> between hybridizati<strong>on</strong> and invasi<strong>on</strong>ability. The genetic heterogeneity <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina hybrids willlead to <str<strong>on</strong>g>the</str<strong>on</strong>g> evoluti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> even more invasive hybridpopulati<strong>on</strong>s.Table 1. Genotype means and percent change between plots with S. virginica "clipped" vs. unclipped, end-<str<strong>on</strong>g>of</str<strong>on</strong>g>-seas<strong>on</strong> shoot densityand aboveground dry biomass.Spartina Shoot Density (#/m 2 ) Dry Biomass (g/m 2 )Genotype Unclipped Clipped % Change Unclipped Clipped % Change2-15 (S. foli) 41 84 104 107 206 931-9 107 235 119* 212 330 562-17 84 117 39 234 243 41-5 67 136 103* 695 1360 96*3-13 77 173 126* 277 426 543-1 74 117 59 464 471 21-17 65 104 60 512 1251 144*2-8 62 139 124* 501 556 113-8 46 142 211* 159 342 115* Significant difference (P < 0.05)-30-
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