<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> SpartinaIMPACT OF INVASIVE HYBRID CORDGRASS (SPARTINA ALTERNIFLORA X FOLIOSA) ONSONG SPARROW AND MARSH WREN POPULATIONS IN SAN FRANCISCO BAY SALTMARSHESJ.C. NORDBY 1,4 , J.T. MCBROOM 2 ,A.N.COHEN 3 AND S.R. BEISSINGER 11Department <str<strong>on</strong>g>of</str<strong>on</strong>g> Envir<strong>on</strong>mental Science, Policy and Management, University <str<strong>on</strong>g>of</str<strong>on</strong>g> California, Berkeley, 151 Hilgard Hall #3110Berkeley, CA, 947202San Francisco Estuary <strong>Invasive</strong> Spartina Project, 2612-A 8 th Street, Berkeley, CA 947103 San Francisco Estuary Institute, 7770 Pardee Lane, 2 nd Floor, Oakland, CA 946214Current address: Institute <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Envir<strong>on</strong>ment and Department <str<strong>on</strong>g>of</str<strong>on</strong>g> Ecology and Evoluti<strong>on</strong>ary Biology, University <str<strong>on</strong>g>of</str<strong>on</strong>g>California, Los Angeles, Los Angeles, CA 90095; nordby@ucla.eduExotic hybrid cordgrass, Spartina alterniflora x foliosa, is altering <str<strong>on</strong>g>the</str<strong>on</strong>g> vegetative structure andcompositi<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 Bay tidal marsh ecosystem, which has multiple impacts <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g>Alameda s<strong>on</strong>g sparrow (Melospiza melodia pusillula), a resident passerine species that is aCalifornia Species <str<strong>on</strong>g>of</str<strong>on</strong>g> Special C<strong>on</strong>cern. These sparrows are affected not <strong>on</strong>ly by <str<strong>on</strong>g>the</str<strong>on</strong>g> altered habitat,but also by <str<strong>on</strong>g>the</str<strong>on</strong>g> occupati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> this habitat by a potential competitor, <str<strong>on</strong>g>the</str<strong>on</strong>g> marsh wren (Cistothoruspalustris). To assess <str<strong>on</strong>g>the</str<strong>on</strong>g> impact <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> S. alterniflora invasi<strong>on</strong> <strong>on</strong> s<strong>on</strong>g sparrow and marsh wrenpopulati<strong>on</strong>s we: 1) located s<strong>on</strong>g sparrow nests to observe nesting habitat preferences and nestsuccess and 2) used focal observati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> color-banded birds to assess <str<strong>on</strong>g>the</str<strong>on</strong>g> vegetati<strong>on</strong> compositi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g>each territory and <str<strong>on</strong>g>the</str<strong>on</strong>g> amount <str<strong>on</strong>g>of</str<strong>on</strong>g> territory overlap between <str<strong>on</strong>g>the</str<strong>on</strong>g> two species. Our findings suggest that<str<strong>on</strong>g>the</str<strong>on</strong>g> changes in salt marsh habitat associated with <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> cordgrass hybrids may favor marshwrens over s<strong>on</strong>g sparrows and could eventually result in a decrease in salt marsh s<strong>on</strong>g sparrowpopulati<strong>on</strong>s.Keywords: Spartina alterniflora, s<strong>on</strong>g sparrow, marsh wren, nest success, territory, competiti<strong>on</strong>,San Francisco BayINTRODUCTIONNative San Francisco Bay tidal salt marshes arecharacterized by broad expanses <str<strong>on</strong>g>of</str<strong>on</strong>g> open tidal mudflats, anarrow mid-marsh z<strong>on</strong>e where Spartina foliosa occurs, and ahigh-marsh z<strong>on</strong>e composed mainly <str<strong>on</strong>g>of</str<strong>on</strong>g> low-growingSarcocornia spp. (formerly Salicornia spp.) with narrowareas <str<strong>on</strong>g>of</str<strong>on</strong>g> Grindelia that line <str<strong>on</strong>g>the</str<strong>on</strong>g> meandering tidal channels.Spartina alterniflora, a cordgrass native to <str<strong>on</strong>g>the</str<strong>on</strong>g> Atlantic andGulf coasts <str<strong>on</strong>g>of</str<strong>on</strong>g> North America, was introduced to SanFrancisco Bay in <str<strong>on</strong>g>the</str<strong>on</strong>g> early 1970s (Ayres et al. 2004). Theexotic cordgrass subsequently hybridized with <str<strong>on</strong>g>the</str<strong>on</strong>g> nativecordgrass, S. foliosa, and <str<strong>on</strong>g>the</str<strong>on</strong>g>se hybrids have spread and nowcover more than 600 hectares (ha) <str<strong>on</strong>g>of</str<strong>on</strong>g> tidal flat and tidalmarsh habitat (Zaremba et al., this volume).The tall, dense exotic Spartina (S. alterniflora and/or<str<strong>on</strong>g>the</str<strong>on</strong>g> hybrid S. alterniflora x foliosa) can grow fur<str<strong>on</strong>g>the</str<strong>on</strong>g>r down<str<strong>on</strong>g>the</str<strong>on</strong>g> tidal gradient than any native tidal marsh plant speciesand so is able to col<strong>on</strong>ize open tidal flats. This exoticcordgrass can also grow much fur<str<strong>on</strong>g>the</str<strong>on</strong>g>r up <str<strong>on</strong>g>the</str<strong>on</strong>g> tidal gradientthan native S. foliosa and thus displaces o<str<strong>on</strong>g>the</str<strong>on</strong>g>r native plantspecies in <str<strong>on</strong>g>the</str<strong>on</strong>g> mid- to high-marsh z<strong>on</strong>es as well (Ayres et al.1999; Nordby pers. obs.).The pr<str<strong>on</strong>g>of</str<strong>on</strong>g>ound changes in habitat structure andcompositi<strong>on</strong> that accompany <str<strong>on</strong>g>the</str<strong>on</strong>g> exotic Spartina invasi<strong>on</strong>(Callaway and Josselyn 1992) will likely have <str<strong>on</strong>g>the</str<strong>on</strong>g> greatestimpact <strong>on</strong> species, such as birds, that are wholly dependent<strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> tidal salt marsh system. The Alameda s<strong>on</strong>g sparrowFig. 1. Alameda s<strong>on</strong>g sparrow (Melospiza melodia pusillula). Photo byJen McBroom.- 197 -
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> SpartinaCriteri<strong>on</strong>) revealed that overall nesting success wasestimated to be 30% lower in areas <str<strong>on</strong>g>of</str<strong>on</strong>g> exotic Spartina than inareas <str<strong>on</strong>g>of</str<strong>on</strong>g> native vegetati<strong>on</strong>. We also found str<strong>on</strong>g evidence <str<strong>on</strong>g>of</str<strong>on</strong>g>marsh wren destructi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> sparrrow eggs, particularly inareas <str<strong>on</strong>g>of</str<strong>on</strong>g> high marsh wren density (Nordby et al. 2009).Fig. 2. Marsh wren (Cistothorus palustris). Photo by Jen McBroom.(Melospiza melodia pusillula) (Fig. 1), a California Species<str<strong>on</strong>g>of</str<strong>on</strong>g> Special C<strong>on</strong>cern, resides entirely within salt marshes inSouth San Francisco Bay. In a native marsh, this sparrow is<str<strong>on</strong>g>the</str<strong>on</strong>g> main resident passerine species and occupies territoriesand nests in Sarcocornia and Grindelia near tidal channels.Ano<str<strong>on</strong>g>the</str<strong>on</strong>g>r passerine species, <str<strong>on</strong>g>the</str<strong>on</strong>g> marsh wren (Cistothoruspalustris) (Fig. 2), that normally occurs in fresh or brackishwater marshes <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> Pacific Coast, has started to occupy <str<strong>on</strong>g>the</str<strong>on</strong>g>newly available exotic Spartina habitat (Nordby and Cohen,pers. obs.). Marsh wrens are very aggressive and will defend<str<strong>on</strong>g>the</str<strong>on</strong>g>ir territories against o<str<strong>on</strong>g>the</str<strong>on</strong>g>r birds, even o<str<strong>on</strong>g>the</str<strong>on</strong>g>r species, bybreaking <str<strong>on</strong>g>the</str<strong>on</strong>g> eggs in nests that are close to <str<strong>on</strong>g>the</str<strong>on</strong>g>ir ownterritories (Picman 1977).To assess <str<strong>on</strong>g>the</str<strong>on</strong>g> impact <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> exotic Spartina invasi<strong>on</strong> <strong>on</strong>s<strong>on</strong>g sparrow and marsh wren populati<strong>on</strong>s in San FranciscoBay we 1) studied sparrow nesting habitat preferences andnest success and also looked for evidence <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> destructi<strong>on</strong><str<strong>on</strong>g>of</str<strong>on</strong>g> s<strong>on</strong>g sparrow eggs by marsh wrens, and 2) assessed <str<strong>on</strong>g>the</str<strong>on</strong>g>vegetati<strong>on</strong> compositi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> sparrow and wren territories aswell as <str<strong>on</strong>g>the</str<strong>on</strong>g> amount <str<strong>on</strong>g>of</str<strong>on</strong>g> territory overlap between <str<strong>on</strong>g>the</str<strong>on</strong>g> twospecies.SONG SPARROW NEST SUCCESSDuring <str<strong>on</strong>g>the</str<strong>on</strong>g> 2002 and 2003 breeding seas<strong>on</strong>s wefollowed and observed <str<strong>on</strong>g>the</str<strong>on</strong>g> fates <str<strong>on</strong>g>of</str<strong>on</strong>g> 351 nests in 45+territories across three study sites (Newark, San Leandro andAlameda). Once <str<strong>on</strong>g>the</str<strong>on</strong>g> fate <str<strong>on</strong>g>of</str<strong>on</strong>g> a nest had been determined(failed due to predati<strong>on</strong>, failed due to tidal flooding, failedfor o<str<strong>on</strong>g>the</str<strong>on</strong>g>r reas<strong>on</strong>s, or successful), we recorded <str<strong>on</strong>g>the</str<strong>on</strong>g> locati<strong>on</strong>and vegetati<strong>on</strong> compositi<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> nest site.We found that s<strong>on</strong>g sparrows did use exotic Spartina asnesting habitat, but <str<strong>on</strong>g>the</str<strong>on</strong>g>se nests were much more likely to faildue to tidal flooding than nests placed in native vegetati<strong>on</strong>.As a result, our model <str<strong>on</strong>g>of</str<strong>on</strong>g> daily nest survival (a generalizedlinear modeling approach using Akaike’s Informati<strong>on</strong>SONG SPARROW AND MARSH WREN TERRITORIESDuring <str<strong>on</strong>g>the</str<strong>on</strong>g> 2003 breeding seas<strong>on</strong> (from March toAugust), we c<strong>on</strong>ducted two to four focal observati<strong>on</strong>s <strong>on</strong>each color-banded male to map <str<strong>on</strong>g>the</str<strong>on</strong>g> territory boundaries for<str<strong>on</strong>g>the</str<strong>on</strong>g> 32 s<strong>on</strong>g sparrows and 16 marsh wrens in <str<strong>on</strong>g>the</str<strong>on</strong>g> twoSpartina-invaded study sites (San Leandro and Alameda).Observers used binoculars, a compass, and a range-finder tomap bird locati<strong>on</strong>s during each <strong>on</strong>e-hour observati<strong>on</strong> period.Observati<strong>on</strong> locati<strong>on</strong>s were marked using a GPS unit, birdlocati<strong>on</strong>s were <str<strong>on</strong>g>the</str<strong>on</strong>g>n calculated and <str<strong>on</strong>g>the</str<strong>on</strong>g>se data were added toan ArcView GIS database. The points were used to c<strong>on</strong>struct100% minimum c<strong>on</strong>vex polyg<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> each territory. Usingcolor-infrared aerial photographs, we identified areas <str<strong>on</strong>g>of</str<strong>on</strong>g>invasive Spartina throughout <str<strong>on</strong>g>the</str<strong>on</strong>g> sites. In ArcView, wecombined <str<strong>on</strong>g>the</str<strong>on</strong>g> layer <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina vegetati<strong>on</strong> with <str<strong>on</strong>g>the</str<strong>on</strong>g> layers <str<strong>on</strong>g>of</str<strong>on</strong>g>marsh wren and s<strong>on</strong>g sparrow territory polyg<strong>on</strong>s. We <str<strong>on</strong>g>the</str<strong>on</strong>g>ndetermined <str<strong>on</strong>g>the</str<strong>on</strong>g> percentage <str<strong>on</strong>g>of</str<strong>on</strong>g> each territory that wascomposed <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina habitat, as well as <str<strong>on</strong>g>the</str<strong>on</strong>g> amount <str<strong>on</strong>g>of</str<strong>on</strong>g>overlap between <str<strong>on</strong>g>the</str<strong>on</strong>g> two species.Although s<strong>on</strong>g sparrows did include some exoticSpartina habitat in <str<strong>on</strong>g>the</str<strong>on</strong>g>ir territories, all but <strong>on</strong>e s<strong>on</strong>g sparrowterritory included some areas <str<strong>on</strong>g>of</str<strong>on</strong>g> native salt marsh habitat.The <strong>on</strong>e territory that was determined from aerialphotographs to be entirely covered by Spartina, actually hada large porti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> native pickleweed vegetati<strong>on</strong> underlying<str<strong>on</strong>g>the</str<strong>on</strong>g> Spartina stand. In c<strong>on</strong>trast, marsh wren territories weremore highly correlated with exotic Spartina habitat andmany territories were exclusively composed <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> exoticcordgrass. We also found that <str<strong>on</strong>g>the</str<strong>on</strong>g>re was little overlapbetween <str<strong>on</strong>g>the</str<strong>on</strong>g> territories <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> two species (Nordby et al., inprep).DISCUSSIONThese results suggest that <str<strong>on</strong>g>the</str<strong>on</strong>g> changes in salt marshhabitat associated with <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> exotic Spartina mayfavor marsh wrens over s<strong>on</strong>g sparrows. While s<strong>on</strong>g sparrowsare occupying and nesting in <str<strong>on</strong>g>the</str<strong>on</strong>g> exotic Spartina, those thatdo so may be at a disadvantage. It is possible that s<strong>on</strong>gsparrows are being drawn to nesting sites in exotic Spartinathat are inappropriate because <str<strong>on</strong>g>the</str<strong>on</strong>g>y are too low in elevati<strong>on</strong>relative to <str<strong>on</strong>g>the</str<strong>on</strong>g> tides. This increase in nest failure due t<str<strong>on</strong>g>of</str<strong>on</strong>g>looding coupled with an apparent increase in interferencecompetiti<strong>on</strong> from marsh wrens may serve to negativelyimpact salt marsh s<strong>on</strong>g sparrow populati<strong>on</strong>s in SanFrancisco Bay.However, we do not yet know whe<str<strong>on</strong>g>the</str<strong>on</strong>g>r exotic Spartinais acting as an ‘ecological trap’ for s<strong>on</strong>g sparrows, where- 198 -
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FORWARD & ACKNOWLEDGEMENTSThe <stro
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CHAPTER ONESpartina Biology
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