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The Impact of Invasive Plants on Tidal-Marsh Vertebrate Species: Common Reed (Phragmites Australis) and Smooth Cordgrass (Spartina Alterniflora) As Case Studies

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The Impact of Invasive Plants on Tidal-Marsh Vertebrate Species: Common Reed (Phragmites Australis) and Smooth Cordgrass (Spartina Alterniflora) As Case Studies Studies in Avian Biology No. 32:229–237 THE IMPACT OF INVASIVE PLANTS ON TIDAL-MARSH VERTEBRATE SPECIES: COMMON REED (PHRAGMITES AUSTRALIS) AND SMOOTH CORDGRASS (SPARTINA ALTERNIFLORA) AS CASE STUDIES GLENN R. GUNTENSPERGEN AND J. CULLY NORDBY Abstract. Large areas of tidal marsh in the contiguous US and the Maritime Provinces of Canada are threatened by invasive plant species. Our understanding of the impact these invasions have on tidal-marsh vertebrates is sparse. In this paper, we focus on two successful invasive plant taxa that have spread outside their native range—common reed (Phragmites australis) and smooth cordgrass (Spartina alternifl ora). A cryptic haplotype of common reed has expanded its range in Atlantic Coast tidal marshes and smooth cordgrass, a native dominant plant of Atlantic Coast low-marsh habitat, has expanded its range and invaded intertidal-marsh habitats of the Pacifi c Coast. The invasions of common reed in Atlantic Coast tidal marshes and smooth cordgrass in Pacifi c Coast tidal marshes appear to have similar impacts. The structure and composition of these habitats has been altered and invasion and dominance by these two taxa can lead to profound changes in geomorphological processes, altering the vertical relief and potentially affecting invertebrate communities and the entire trophic structure of these systems. Few studies have documented impacts of invasive plant taxa on tidal-marsh vertebrate species in North America. However, habitat specialists that are already con- sidered threatened or endangered are most likely to be affected. Extensive experimental studies are needed to examine the direct impact of invasive plant species on native vertebrate species. Careful monitoring of sites during the initial stages of plant invasion and tracking ecosystem changes through time are essential. Since tidal marshes are the foci for invasion by numerous species, we also need to understand the indirect impacts of invasion of these habitats on the vertebrate community. We also suggest the initiation of studies to determine if vertebrate species can compensate behaviorally for alterations in their habitat caused by invasive plant species, as well as the potential for adaptation via rapid evolution. Finally, we urge natural-resource managers to consider the impact various invasive plant control strategies will have on native vertebrate communities. Key Words: food webs, geomorphology, invasive plants, marsh birds, North America, Phragmites, Spartina alternifl ora, saltmarsh, vertebrates. EL IMPACTO DE PLANTAS INVASORAS EN ESPECIES DE VERTEBRADOS EN MARISMA DE MAREA: EL CARRIZO (PHRAGMITES AUSTRALIS) Y EL PASTO (SPARTINA ALTERNIFLORA) COMO CASOS DE ESTUDIO Resumen. Grandes áreas de marisma de marea a lo largo EU así como y en las Provincias de Marisma de Canadá, se encuentran amenazadas por especies de plantas invasoras. Nuestro entendimiento acerca del impacto que estas especies tienen en los vertebrados de marismas de marea es escaso. En este artículo nos enfocamos en dos taxa de especies de plantas exitosas que se han dispersado fuera de su rango nativo—el carrizo (Phragmites australis) y el pasto (Spartina alternifl ora). Un haplotipo críptico de carrizo ha expandido su rango en las marismas de marea en la Costa del Atlántico. El pasto (Spartina alternifl ora), nativa y dominante del hábitat de marisma baja de la Costa del Atlántico, ha expandido su rango e invadido habitats de marisma intermareal en la Costa Pacífi co. Las invasiones del carrizo en las marismas de marea de la Costa Atlántica y el pasto en marismas de marea de la Costa del Pacífi co parece que tienen impactos similares. La estructura y composición de estos habitats ha sido alterada y la invasión y dominancia por estas dos taxa, pueden derivar en cambios profundos en los procesos geomorfológicos, alterando la mitigación vertical y pueden potencialmente afectar las comunidades de invertebrados y toda la estructura trófi ca de estos ecosistemas. Pocos estudios han documentado impactos de taxa de plantas invasoras en especies de vertebrados de marisma de marea en Norte América. Sin embargo, especialistas del hábitat, los cuales ya están considerados como en peligro, son los que están siendo más afectados. Se necesitan estudios extensivos experimentales para examinar el impacto directo de especies de plantas invasoras en especies nativas de vertebrados. El monitoreo cauteloso de los sitios durante los estados iniciales de la invasión de plantas y el rastreo de los cambios del ecosistema en el tiempo son esenciales. Debido a que las marismas de marea son el foci para la invasión por numerosas especies, también necesitamos los impactos indirectos de invasión de estos habitats en la comunidad de vertebrados. También sugerimos el inicio de estudios para determinar si especies de vertebrados pueden compensarse en términos de comportamiento por las alteraciones en su hábitat causado por especies de plantas invasoras, como también el potencial para la adaptación vía evolución rápida. Finalmente, recomendamos a los manejadores de recursos 229 230 STUDIES IN AVIAN BIOLOGY NO. 32 naturales a que consideren el impacto que tendrán las estrategias de control de varias plantas invasoras en comunidades nativas de vertebrados. Invasive plant taxa are profoundly chang- the Hudson River north to the St. Lawrence ing North American saltmarshes, but this is estuary, the coastal plain of the United States not an isolated phenomenon. The introduction from New Jersey south along the southeast- of non-indigenous plants in diverse habitats ern US Atlantic Coast to the northern Gulf of represents some of the most dramatic examples Mexico, and the western US along the Pacifi c of biological invasions. Their impact on natural coast (Dame et al. 2000, Emmett et al. 2000, habitats, and the biodiversity of those habitats, Roman et al. 2000). In this chapter, we are inter- is a pervasive threat and one of the most daunt- ested in two areas—the Atlantic Coast, with an ing ecological challenges facing twenty-fi rst emphasis on the northeastern Atlantic region, century natural-resource managers. and the Pacifi c Coast. Considerable attention has been devoted Northeast-coast saltmarshes are formed to understanding the attributes of successful largely by reworked marine sediments and in invaders and the characteristics of invaded situ peat formation. These marshes are largely regions and habitats, as well as to documenting limited to small, narrow fringing systems the patterns and history of invasion (Mooney because the physiography of the region and and Drake 1986, Drake et al. 1989, Cronk and broad expanses of rocky coast limit their areal Fuller 1995, Pysek et al. 1995). The consequences extent (Nixon 1982). Farther south, more exten- of invasive taxa on biological communities and sive saltmarshes occur in the drowned valley ecosystem processes have been documented estuaries of Delaware Bay and Chesapeake Bay more recently (D’Antonio and Vitousek 1992, (Teal 1986). Pacifi c Coast saltmarshes occur in Mack et al. 2000). MacDonald et al. (1989) calcu- a geologically young region structured by tec- lated that of the 941 vertebrate species thought tonic and volcanic forces (Emmett et al. 2000). to be in danger of extinction worldwide, 18.4% Because of the rocky and unfeatured wave- are threatened in some way by introduced spe- dominated shoreline along much of the Pacifi c cies. In North America, they calculated that Coast, extensive areas of saltmarsh are restricted >13.3% of the native avifauna is threatened by to large estuaries such as those associated with invasive species. In another study focusing on the San Francisco Bay and the Columbia River threats to biodiversity in the US, Wilcove et al. or behind sheltering bay-mouth bars. (1998) found that 49% of all imperiled species Strong physical gradients of salinity and (plants and animals) were threatened by inva- tidal inundation contribute to the characteris- sive species. tic patterns of tidal-height zonation. The two The study of biological invasions has only main marsh zones include low marsh and recently focused on coastal and estuarine high marsh. Low marsh is lower in elevation habitats (Grosholz 2002). Large numbers of relative to mean low water and is regularly non-indigenous species have been identifi ed fl ooded by tides. High marsh occupies the in U.S. coastal estuaries >200 non-indigenous higher elevations in the intertidal zone, and is species from San Francisco Bay alone (Cohen less infl uenced by tidal forces. and Carlton 1998). Most research has concen- The organization of tidal-marsh vegetation trated on non-native aquatic invasive species communities varies in the different regions. including crustaceans, clams, crabs, and hydro- A short-statured grass, saltmeadow cordgrass zoans (Cordell and Morrison 1996, Crooks 1998, (Spartina patens), dominates the high marsh Bagley and Geller 2000, Byers 2000). In contrast, along the northeast coast—often intermixed efforts to examine the ecological effects of non- with the short form of smooth cordgrass and native emergent wetland plant taxa in salt- black needlerush (Juncus gerardi) at the upland marshes have lagged behind (except Weinstein border of the high marsh. Smooth cordgrass et al. 2003). In this chapter, we focus on two of persists as a dominant species in low-marshes, the more problematic invasive marsh-plant taxa in this region reaching heights as tall as 1.25–2 m in North American saltmarshes—common reed (Teal 1986). Open tidal mudfl ats characterize (Phragmites australis) and smooth cordgrass the lower intertidal zone of Pacifi c Coast estu- (Spartina alternifl ora). aries. The mid-intertidal zone is dominated by California cordgrass (Spartina foliosa), which ATLANTIC AND PACIFIC TIDAL SALTMARSHES forms a narrow band of vegetation along the outer edges of the native terrestrial vegetative In the contiguous US and Maritime Provinces zone and along tidal channels (Mahall and Park of Canada saltmarshes occur in three distinct 1976, Ayers et al. 2003). California cordgrass’s regions: the Northeastern Atlantic Coast from range extends from Baja California north to IMPACT OF INVASIVES—Guntenspergen and Nordby 231 Humboldt Bay (Josselyn 1983).
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