<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> SpartinaCONTRASTING EFFECTS OF SPARTINA FOLIOSA AND HYBRID SPARTINA ON BENTHICINVERTEBRATESE.D. BRUSATI 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, One Shields Ave., Davis, CA, USA 956162 Present address: California <strong>Invasive</strong> Plant Council, 1442-A Walnut St. #462, Berkeley, CA USA 94709;edbrusati@cal-ipc.orgIn San Francisco Bay, California, mudflats and native California cordgrass (Spartina foliosa)marshes are being invaded by a hybrid cordgrass formed by hybridizati<strong>on</strong> between S. foliosa andintroduced S. alterniflora. We investigated differences in vegetati<strong>on</strong> and sediment structure, benthicinfauna, and food webs within native and invaded Spartina marshes between San Francisco Bay andBodega Bay, California. The greatest impact <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrid Spartina in San Francisco Bay appears to beits alterati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> habitat structure ra<str<strong>on</strong>g>the</str<strong>on</strong>g>r than food webs. Habitat structure differed significantlybetween native and hybrid Spartina. Hybrid Spartina produced greater biomass both above andbelow ground, and taller stem heights. Spartina foliosa c<strong>on</strong>tained significantly higher densities andbiomass <str<strong>on</strong>g>of</str<strong>on</strong>g> infaunal organisms in benthic cores than did mudflats, while densities and biomass <str<strong>on</strong>g>of</str<strong>on</strong>g>infauna in hybrid Spartina were lower than, or did not differ from, mudflats. Stable isotopes <str<strong>on</strong>g>of</str<strong>on</strong>g>carb<strong>on</strong> and nitrogen were used to examine whe<str<strong>on</strong>g>the</str<strong>on</strong>g>r macr<str<strong>on</strong>g>of</str<strong>on</strong>g>aunal food webs differ between native orhybrid Spartina and mudflats. Some c<strong>on</strong>sumers collected within Spartina showed evidence <str<strong>on</strong>g>of</str<strong>on</strong>g> a shiftin carb<strong>on</strong> isotope ratios indicating a possible increase in Spartina c<strong>on</strong>sumpti<strong>on</strong> within vegetati<strong>on</strong>;however, <str<strong>on</strong>g>the</str<strong>on</strong>g> pattern was not c<strong>on</strong>sistent across species and sites. Due to <str<strong>on</strong>g>the</str<strong>on</strong>g> differences in <str<strong>on</strong>g>the</str<strong>on</strong>g>ireffects <strong>on</strong> infauna, hybrid Spartina and S. foliosa should not be c<strong>on</strong>sidered equivalent for marshrestorati<strong>on</strong> projects.Keywords: hybrid Spartina, Spartina foliosa, infauna, food webs, CaliforniaINTRODUCTIONIn San Francisco Bay, hybrids formed between nativeCalifornia cordgrass (S. foliosa) and introduced S.alterniflora accrete sediment, increase elevati<strong>on</strong> compared tosurrounding mudflats, and significantly reduce light levelsunder <str<strong>on</strong>g>the</str<strong>on</strong>g>ir canopies (Neira et al. 2005). Based <strong>on</strong> tidallevels, hybrid Spartina would also be able to fill large areas<str<strong>on</strong>g>of</str<strong>on</strong>g> shallow outer coast bays if it establishes populati<strong>on</strong>s <str<strong>on</strong>g>the</str<strong>on</strong>g>re(Daehler and Str<strong>on</strong>g 1997). The spread <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrid Spartinathreatens flood c<strong>on</strong>trol channels and habitat for migratingshorebirds, and may impact invertebrates that are food forbirds and fishes.Spartina cordgrasses may ei<str<strong>on</strong>g>the</str<strong>on</strong>g>r facilitate or inhibitinfauna depending <strong>on</strong> locati<strong>on</strong> (e.g., Capehart and Hackney1989, Netto and Lana 1999). The invasi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> hybridSpartina provides an opportunity to examine howdifferences in structure between two closely-relatedecosystem engineers affect infaunal communities.Understanding differences between S. foliosa and hybridSpartina may help managers predict <str<strong>on</strong>g>the</str<strong>on</strong>g> impacts <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>c<strong>on</strong>tinued spread <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrid Spartina.This study examined whe<str<strong>on</strong>g>the</str<strong>on</strong>g>r hybrid Spartina isecologically equivalent to native S. foliosa in its impacts <strong>on</strong>infaunal and epifaunal invertebrates. We investigated twohypo<str<strong>on</strong>g>the</str<strong>on</strong>g>ses. First, we predicted that differences in structurebetween S. foliosa and hybrid Spartina will be reflected indifferences in infaunal density, biomass, and tax<strong>on</strong>omiccompositi<strong>on</strong>. Sec<strong>on</strong>d, we predicted that, due to <str<strong>on</strong>g>the</str<strong>on</strong>g> greateraboveground biomass produced by hybrid Spartina,organisms living within <str<strong>on</strong>g>the</str<strong>on</strong>g> hybrid will show greater use <str<strong>on</strong>g>of</str<strong>on</strong>g>hybrid as a food source than those in S. foliosa or mudflats,based <strong>on</strong> stable isotopes <str<strong>on</strong>g>of</str<strong>on</strong>g> carb<strong>on</strong> and nitrogen.METHODSStudy sites included five S. foliosa marshes and twohybrid Spartina marshes in nor<str<strong>on</strong>g>the</str<strong>on</strong>g>rn California. Spartinafoliosa sites included: China Camp State Park (38° 0.37’N,122° 28.66’W) <strong>on</strong> San Pablo Bay; Bolinas Lago<strong>on</strong> (38°19.22’N, 122° 41.73’W); Shields Marsh (38° 5.35’N, 122°50.44’W) and Tom’s Point (38° 13.21’N, 122° 56.86’W) <strong>on</strong>Tomales Bay; Drakes Estero (38° 5.36’N, 122° 55.86’W).Hybrid marshes were located <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> eastern side <str<strong>on</strong>g>of</str<strong>on</strong>g> SanFrancisco Bay in San Lorenzo (Roberts Landing, 37°40.22’N 122° 09.70’W) and Alameda (Elsie Roemer BirdSanctuary, 37° 45.58’N 122° 28.80’W). The San Lorenzosite is unique in that it c<strong>on</strong>tains discrete patches <str<strong>on</strong>g>of</str<strong>on</strong>g> both S.foliosa and hybrid Spartina (genotypes c<strong>on</strong>firmed by D.Ayres, University <str<strong>on</strong>g>of</str<strong>on</strong>g> California, Davis, pers. comm.).- 161 -
Chapter 3: Ecosystem Effects <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>Invasive</strong> Spartina<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> SpartinaWe established ten study quadrats within Spartina ateach site, paired with ten <strong>on</strong> mudflats. These quadrats wereused for vegetati<strong>on</strong>, sediment, and infauna sampling. Fordetails <str<strong>on</strong>g>of</str<strong>on</strong>g> methods, see Brusati and Grosholz (2006). Wemeasured stem heights and densities, aboveground biomass,and belowground biomass at each site, as well as sedimentcharacteristics such as organic matter c<strong>on</strong>tent, bulk density,sediment porewater salinity, and oxidati<strong>on</strong>-reducti<strong>on</strong>potential. Infauna cores (5 centimeters [cm] deep x 5 cmdiameter) were collected in winter and summer 2001-03.Cores were preserved in 8% formalin and organisms werecounted and weighed in <str<strong>on</strong>g>the</str<strong>on</strong>g> laboratory. We used t-tests <strong>on</strong>transformed data to compare results between Spartina andmudflats.To analyze <str<strong>on</strong>g>the</str<strong>on</strong>g> effects <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina <strong>on</strong> invertebrate foodwebs, we collected species <str<strong>on</strong>g>of</str<strong>on</strong>g> infaunal and epifaunalinvertebrates in S. foliosa, hybrid Spartina, and mudflats.To understand how <str<strong>on</strong>g>the</str<strong>on</strong>g>se plants are incorporated into <str<strong>on</strong>g>the</str<strong>on</strong>g>invertebrate food webs we analyzed organisms for naturallyoccuring abundances <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> stable isotope ratios 13 C and 15 N. The carb<strong>on</strong> signal reflects a weighted average <str<strong>on</strong>g>of</str<strong>on</strong>g> foodsources, with Spartina 13 C = -14 ‰, which is significantlymore enriched than o<str<strong>on</strong>g>the</str<strong>on</strong>g>r carb<strong>on</strong> sources in this habitat(Cloern et al. 2002). We predict that if hybrid Spartina isentering <str<strong>on</strong>g>the</str<strong>on</strong>g> food web, c<strong>on</strong>sumers collected within invadedareas will have a str<strong>on</strong>ger Spartina signal than those frommudflats, with c<strong>on</strong>sumers from S. foliosa showing anintermediate signal.RESULTSHybrid Spartina differs from S. foliosa in its effects <strong>on</strong>habitat structure, infaunal communities, and food webs.Hybrid Spartina produces more biomass, and <str<strong>on</strong>g>the</str<strong>on</strong>g>refore moredense structure <strong>on</strong> mudflats, than S. foliosa both above andbelow ground (Fig. 1). Alameda, where <str<strong>on</strong>g>the</str<strong>on</strong>g> invasi<strong>on</strong> is 30years old, has greater belowground biomass than SanLorenzo, which was invaded in <str<strong>on</strong>g>the</str<strong>on</strong>g> 1990s. We found fewc<strong>on</strong>sistent differences between sediment characteristics <str<strong>on</strong>g>of</str<strong>on</strong>g>hybrid or S. foliosa and mudflats. Spartina foliosa generallyhad significantly higher infaunal densities and biomass thanadjacent mudflats. Winter data are presented here, butsummer samples showed similar patterns. In c<strong>on</strong>trast, hybridSpartina sediments never c<strong>on</strong>tained significantly greaterdensities or biomass than mudflats (Fig. 2, numbers abovebars are p-values, n = 10).Food web analysis also shows differences between S.foliosa and hybrid Spartina. Stable isotope results indicated thatsome species show evidence <str<strong>on</strong>g>of</str<strong>on</strong>g> a slight shift in 13 C towardSpartina for individuals collected within S. foliosa. (Fig. 3) Forexample, shore crabs (Hemigrapsus oreg<strong>on</strong>ensis) and Europeangreen crabs (Carcinus maenas) living within S. foliosa at ChinaCamp, and H. oreg<strong>on</strong>ensis in S. foliosa at Drakes Estero hadisotopic signatures more similar to S. foliosa than thosecollected <strong>on</strong> mudflats. At Alameda, <str<strong>on</strong>g>the</str<strong>on</strong>g>re was no significantdifference in isotopic signatures <str<strong>on</strong>g>of</str<strong>on</strong>g> C. maenas or Atlantic oysterdrills (Urosalpinx cinerea) from hybrid Spartina and mudflats.When results from all sites are compared, hybrid Spartina doesnot appear to produce a str<strong>on</strong>ger shift in isotope signatures thanS. foliosa (Brusati and Grosholz 2009).DISCUSSIONOur results show that hybrid Spartina is not ecologicallyequivalent to <str<strong>on</strong>g>the</str<strong>on</strong>g> native cordgrass. While both S. foliosa andhybrid Spartina modify estuarine habitat, <str<strong>on</strong>g>the</str<strong>on</strong>g> magnitude <str<strong>on</strong>g>of</str<strong>on</strong>g><str<strong>on</strong>g>the</str<strong>on</strong>g> changes produced by hybrid Spartina are greater,resulting in qualitative differences in <str<strong>on</strong>g>the</str<strong>on</strong>g> infauna. Thestr<strong>on</strong>gest differences were seen in <str<strong>on</strong>g>the</str<strong>on</strong>g> greater height andaboveground biomass <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrid Spartina compared to S.foliosa. The greater aboveground biomass <str<strong>on</strong>g>of</str<strong>on</strong>g> hybrid Spartinais more than would be expected from differences in stemheight and density between <str<strong>on</strong>g>the</str<strong>on</strong>g> two species; it also reflects<str<strong>on</strong>g>the</str<strong>on</strong>g> fact that <str<strong>on</strong>g>the</str<strong>on</strong>g> hybrid’s stems are c<strong>on</strong>siderably thicker thanS. foliosa’s. Native S. foliosa faciliates infauna, possibly bystabilizing substrate or providing attachment sites for tubebuildingorganisms. In c<strong>on</strong>trast, <str<strong>on</strong>g>the</str<strong>on</strong>g> dense roots andAboveground Biomass (g/m 2 )25002000150010005000Outer coastNodataSan FranciscoEstuaryBH BL DE SH TP CC SLF SLH AL---------------Spartina foliosa ----------------------- Hybrid2001 2002 2003Individuals/m 27000060000500004000030000200001000000.005Outer Coast0.2630.0020.000San FranciscoEstuary0.7240.114BL DE SM TP CC SL ALMudflat S. foliosa Hybrid0.001Fig. 1. Aboveground biomass <str<strong>on</strong>g>of</str<strong>on</strong>g> S. foliosa and hybrid Spartina 2001-03 (n= 10). Site abbreviati<strong>on</strong>s: BH = Bodega Harbor, BL = Bolinas Lago<strong>on</strong>, DE= Drake’s Estero, SH = Shields March, TP = Tom’s Pt., CC = China Camp,SLF = Robert’s Landing S. foliosa, SLH = San Lorenzo hybrid, AL =Alameda.Fig. 2. Infaunal densities in S. foliosa exceed those in mudflats, whilehybrid Spartina never c<strong>on</strong>tained greater densities than mudflats (numbersabove bars are p-values, n = 10, winter 2001 data shown).- 162 -
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