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Proceedings of the Third International Conference on Invasive ...

Proceedings of the Third International Conference on Invasive ...

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<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 3: Ecosystem Effects <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>Invasive</strong> SpartinaSPARTINA INVASION CHANGES INTERTIDAL ECOSYSTEM METABOLISM IN SANFRANCISCO BAYA.C. TYLER 1,2 AND E.D. GROSHOLZ 11 Department <str<strong>on</strong>g>of</str<strong>on</strong>g> Envir<strong>on</strong>mental Science and Policy, University <str<strong>on</strong>g>of</str<strong>on</strong>g> California, Davis, CA 95616;annachristinatyler@gmail.com2 Current address: School <str<strong>on</strong>g>of</str<strong>on</strong>g> Biological and Medical Sciences, Rochester Institute <str<strong>on</strong>g>of</str<strong>on</strong>g> Technology, Rochester, NY 14623In San Francisco Bay, Atlantic smooth cordgrass, Spartina alterniflora, and its hybrids haveinvaded unvegetated mudflats and native marshes formerly dominated by Sarcocorniapacifica and Spartina foliosa. This recent, rapid invasi<strong>on</strong> has dramatically changedecosystem processes and food web structure. We measured sediment fluxes <str<strong>on</strong>g>of</str<strong>on</strong>g> carb<strong>on</strong>dioxide (CO 2 ) to determine microalgal gross primary producti<strong>on</strong> (GPP), sediment respirati<strong>on</strong>and net sediment metabolism in native intertidal areas (mudflats, S. pacifica and S. foliosa)and in adjacent areas invaded by hybrid Spartina. Sediment microalgal GPP and microalgalchlorophyll a were substantially higher in all native habitats than in <str<strong>on</strong>g>the</str<strong>on</strong>g> hybrid. At <str<strong>on</strong>g>the</str<strong>on</strong>g> sametime, sediment respirati<strong>on</strong> rates were generally higher in native vegetati<strong>on</strong> than in hybriddominatedhabitats, but substantially lower <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> mudflats than elsewhere. Net sedimentmetabolism switched from autotrophy to heterotrophy following invasi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> mudflats.However, <str<strong>on</strong>g>the</str<strong>on</strong>g> higher respirati<strong>on</strong> rate in native vegetati<strong>on</strong>, especially S. pacifica, relative tohybrid areas, suggests slower decompositi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrid detritus. This result is corroboratedby litterbag decompositi<strong>on</strong> rates and indicates a build-up and/or export <str<strong>on</strong>g>of</str<strong>on</strong>g> refractory organicmatter following hybrid invasi<strong>on</strong>. The switch from a microalgal-dominated system to arefractory detritus-dominated system has clear implicati<strong>on</strong>s for support <str<strong>on</strong>g>of</str<strong>on</strong>g> higher trophiclevels within <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.Keywords: ecosystem metabolism, carb<strong>on</strong> cycling, Spartina alterniflora, Sarcocornia sp.,Spartina foliosa, benthic microalgaeINTRODUCTIONOne <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> most serious threats to natural ecosystemsand <str<strong>on</strong>g>the</str<strong>on</strong>g> maintenance <str<strong>on</strong>g>of</str<strong>on</strong>g> ecosystem services is <str<strong>on</strong>g>the</str<strong>on</strong>g> invasi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g>n<strong>on</strong>-native plant species (Drake et al. 1989; Vitousek et al.1997). These threats may include <str<strong>on</strong>g>the</str<strong>on</strong>g> extincti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> nativespecies, loss <str<strong>on</strong>g>of</str<strong>on</strong>g> functi<strong>on</strong>al native diversity, changes innutrient cycling and organic matter storage and loss <str<strong>on</strong>g>of</str<strong>on</strong>g>habitat. One <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> most dramatic invasi<strong>on</strong>s in coastalsystems in western North America has been <str<strong>on</strong>g>the</str<strong>on</strong>g> invasi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g>Spartina alterniflora (smooth cordgrass), a native <str<strong>on</strong>g>of</str<strong>on</strong>g> westernAtlantic marshes. This invasi<strong>on</strong> has fostered large-scalealterati<strong>on</strong>s in ecosystem processes in commercially andecologically important estuaries in both California andWashingt<strong>on</strong>.S. alterniflora was first introduced into San FranciscoBay by <str<strong>on</strong>g>the</str<strong>on</strong>g> U.S. Army Corps <str<strong>on</strong>g>of</str<strong>on</strong>g> Engineers in 1973 for marshrestorati<strong>on</strong> (Faber 2000). Subsequent hybridizati<strong>on</strong> with <str<strong>on</strong>g>the</str<strong>on</strong>g>native cordgrass, Spartina foliosa, resulted in a highlysuccessful hybrid populati<strong>on</strong> (henceforth hybrid Spartina)that has col<strong>on</strong>ized ~500 acres, in south and central 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> Bay (Daehler and Str<strong>on</strong>g 1997; Ayres et al. 2004).Hybrid Spartina occupies a wide tidal range that includeshistorically unvegetated mudflats and native marshes <strong>on</strong>cedominated by S. foliosa and Sarcocornia pacifica (Callawayand Josselyn 1992).The Spartina invasi<strong>on</strong> has greatly altered <str<strong>on</strong>g>the</str<strong>on</strong>g> cycling <str<strong>on</strong>g>of</str<strong>on</strong>g>carb<strong>on</strong> (C) in <str<strong>on</strong>g>the</str<strong>on</strong>g> intertidal z<strong>on</strong>e. Hybrid Spartina formsdense cl<strong>on</strong>es, with an average aboveground biomass <str<strong>on</strong>g>of</str<strong>on</strong>g> 1.8 ±0.3 kilograms per square meter (kg m -2 ) 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>growing seas<strong>on</strong> (fall; Tyler et al. 2007). Much <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>aboveground porti<strong>on</strong> senesces during <str<strong>on</strong>g>the</str<strong>on</strong>g> winter, but <str<strong>on</strong>g>the</str<strong>on</strong>g>substantial belowground roots and rhizomes (mean = 4.2 ±0.8 kg m -2 ) persist year-round (Tyler et al. 2007). Thecarb<strong>on</strong> to nitrogen (C:N) ratio <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina is higher than <str<strong>on</strong>g>the</str<strong>on</strong>g>native vegetati<strong>on</strong>, particularly when compared tomicroalgae, and <str<strong>on</strong>g>the</str<strong>on</strong>g> slow decompositi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> this refractorydetritus has resulted in substantial deposits <str<strong>on</strong>g>of</str<strong>on</strong>g> belowgroundorganic matter (Neira et al. 2005). We do not know howoverall ecosystem producti<strong>on</strong> and respirati<strong>on</strong> are changed asa result <str<strong>on</strong>g>of</str<strong>on</strong>g> this invasi<strong>on</strong>, but this is clearly important inunderstanding <str<strong>on</strong>g>the</str<strong>on</strong>g> overall impact <str<strong>on</strong>g>of</str<strong>on</strong>g> invasi<strong>on</strong> <strong>on</strong> estuarineecosystem functi<strong>on</strong>.The lower productivity and low stature <str<strong>on</strong>g>of</str<strong>on</strong>g> nativesou<str<strong>on</strong>g>the</str<strong>on</strong>g>rn California salt marsh plants, which results in morelight reaching <str<strong>on</strong>g>the</str<strong>on</strong>g> sediment surface, may promote microalgal- 135 -

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