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Multiple-Scale Habitat Relationships of Tidal-Marsh Breeding Birds in the San Francisco Bay Estuary

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Multiple-Scale Habitat Relationships of Tidal-Marsh Breeding Birds in the San Francisco Bay Estuary Studies in Avian Biology No. 32:247–269 MULTIPLE-SCALE HABITAT RELATIONSHIPS OF TIDAL-MARSH BREEDING BIRDS IN THE SAN FRANCISCO BAY ESTUARY HILDIE SPAUTZ, NADAV NUR, DIANA STRALBERG, AND YVONNE CHAN Abstract. We modeled the abundance or probability of occurrence of several tidal-marsh-dependent birds found in the San Francisco Bay estuary—the San Pablo Song Sparrow (Melospiza melodia samuelis), Alameda Song Sparrow (M. m. pusillula), Suisun Song Sparrow (M. m. maxillaris), Salt Marsh Common Yellowthroat (Geothlypis trichas sinuosa), California Black Rail (Laterallus jamaicensis coturniculus), and Marsh Wren (Cistothorus palustris)—based on marsh characteristics at several scales. Local habitat variables included vegetation type, structure, and height, and tidal-channel characteristics. Landscape variables included marsh size and confi guration, distance to edge, and type of surrounding land use. For each species considered, both landscape and local habitat factors were signifi cant predictors in multi-variable, multi-scale, linear or logistic regression models. While the best models contained both local and landscape variables, all four bird species were also well predicted by local habitat or land- scape variables alone. Predictor variables differed by species, but each species responded strongly to vegetation composition (specifi c plant species) as well as the overall structure (height or complexity) of the vegetation. Scale effects also differed by species. For Song Sparrows, land-use variables were most important at a relatively small spatial scale (500 m) while for Marsh Wrens and Common Yellowthroats they were important at the largest scale examined (2,000 m). Certain elements of vegetation type and structure, as well as marsh size and confi guration (perimeter to area ratio) and surrounding land use, were important across several species, suggesting a suite of habitat and landscape characteristics that may be useful in identifying sites important to multiple bird species. Key Words: Cistothorus palustris, Geothlypis trichas sinuosa, habitat selection, Laterallus jamaicensis cotur- niculus, Melospiza melodia, San Francisco Bay, tidal marsh. RELACIONES DE HABITAT A ESCALAS MULTIPLES DE AVES REPRODUCTORAS DE MARISMA DE MAREA EN EL ESTUARIO DE LA BAHÍA DE SAN FRANCISCO Resumen. Modelamos la abundancia de la probabilidad de ocurrencia de varias especies de aves dependientes de marisma de marea, encontradas en el estuario de la Bahía de San Francisco—el Gorrión Cantor de San Pablo (Melospiza melodia samuelis), el Gorrión Cantor de Alameda (M. m. pusil- lula), el Gorrión Cantor Suisun (M. m. maxillaris), la Mascarita Común de Marisma Salado (Geothlypis trichas sinuosa), la Polluela Negra de California (Laterallus jamaicensis coturniculus), y el Chivirin Pantanero (Cistothorus palustris)—basados en las características de la marisma a diferentes escalas. Las variables locales incluyeron el tipo de vegetación, estructura y altura, y las características del canal de la marea. Variables del paisaje incluyeron el tamaño de la marisma y su confi guración, la distancia a la orilla, y tipo de uso del suelo de los alrededores. Para cada especie considerada, tanto el paisaje como los factores locales del hábitat fueron vaticinadores signifi cativos en los modelos de multi-variable, multi-escala, linear o de regresión logística. Mientras que los mejores modelos con- tenían tanto variables locales como de paisaje, las cuatro especies fueron también bien pronosticadas por el hábitat local o las variables de paisaje solas. Las variables de predicción se diferenciaron por especies, pero cada especie respondió fuertemente a la composición de la vegetación (especies de planta específi cas) como también a la estructura total (altura o complejidad) de la vegetación. Efectos de escala también difi rieron por las especies. Para los Gorriones Cantores, las variables del uso del suelo fueron más importantes a una escala espacial relativamente pequeña (500 m), mientras para los Chivirines Pantaneros y las Mascaritas Comunes de Marisma Salada fueron más importantes a la escala mayor examinada (2,000 m). Ciertos elementos del tipo y de la estructura de la vegetación, como también el tamaño y la confi guración de la marisma (perímetro al radio del área) y el uso del suelo de los alrededores, fueron importantes a través de algunas especies, sugiriendo un juego de características del hábitat y el paisaje que quizás sea utilizado para identifi car sitios importantes para múltiples especies de aves. Tidal marsh, formerly the dominant habitat development (Goals Project 1999). In addi- type in the San Francisco Bay estuary (here- tion, many remaining tidal marshes have after the estuary), has been reduced to <20% been hydrologically altered and subdivided of its original extent as a result of human by levees, mosquito-control ditches, board- activities, such as diking, dredging, and urban walks, and power lines. Many have also been 247 248 STUDIES IN AVIAN BIOLOGY NO. 32 degraded by contaminants, invasive species, 21 San Francisco Bay estuary tidal marshes, and recreational use (Takekawa et al., chapter adding new sites each year to result in a total 11, this volume). This habitat loss and degrada- of 79 marshes surveyed at least once between tion has adversely affected a unique assemblage 1996 and 2003. This comprehensive dataset of marsh-dependent plants, animals, and inver- provides a unique opportunity to examine tebrates, many of which are specifi cally adapted regional distribution and abundance patterns to the range of salinity and tidal regimes in the and, most importantly, to assess the effects estuary’s marshes. of local habitat characteristics, landscape Tidal-marsh passerine birds, including three composition, and habitat fragmentation on endemic subspecies of Song Sparrow (San these distribution and abundance patterns. Pablo Song Sparrow [Melospiza melodia samu- Knowledge of specifi c habitat requirements of elis], Alameda Song Sparrow [M. m. pusillula], these tidal-marsh birds will improve the abil- and Suisun Song Sparrow [M. m. maxillaries] ity of land managers and wildlife agencies to hereafter referred to as Song Sparrows or tidal- plan restoration, management, and acquisition marsh Song Sparrows), the endemic Salt Marsh activities. Common Yellowthroat (Geothlypis trichas sinu- All bird species display some degree of osa), and the Marsh Wren (Cistothorus palustris), specifi city in terms of the habitat types in have experienced a severe habitat loss, and which they choose to set up territories, for- have been restricted in many areas to isolated age, seek shelter, and breed; habitat relation- and degraded marsh fragments with extensive ship models seek to quantify and clarify these urban upland edges. All but the Marsh Wren apparent preferences (Cody 1985). Birds tend are considered species of special concern by to respond to particular characteristics of veg- the state of California. The California Black Rail etation structure and patchiness, often at sev- (Laterallus jamaicensis coturniculus), a state of eral scales (Rotenberry and Wiens 1980, Wiens California threatened species and a federal spe- and Rotenberry 1981, Saab 1999). Habitat char- cies of management concern, is a tidal-marsh- acteristics found to be important for wetland dependent species that is now absent from birds include various aspects of vegetation many estuary marshes, and its small population structure and density (Collins and Resh 1985, size raises concerns about its long-term persis- Leonard and Picman 1987, Weller 1994, Benoit tence in the estuary (Evens et al. 1991, Nur et al. and Askins 1999, Whitt et al. 1999, Poulin et 1997). All of these species merit special attention al. 2002), water depth and cover (Leonard due to their limited distributions and relatively and Picman 1987, Craig and Beal 1992), and small population sizes, but they may also serve tidal-channel characteristics (Collins and Resh as habitat indicators for other tidal-marsh- 1985). However, many bird species may also dependent plant and animal species, several of respond to the landscape context of a habitat which have state and/or federal threatened or patch, as well as its size and shape. Numerous endangered status. studies over a range of habitat types have dem- Important earlier studies of the three focal onstrated a signifi cant effect of surrounding songbird species in the San Francisco Bay estu- landscape at various scales on species richness, ary, primarily concerning the Song Sparrow relative abundance, and nest success of breed- (Johnson 1956a, b; Collins and Resh 1985, ing passerines (Flather and Sauer 1996, Bolger Marshall and Dedrick 1994), were based on et al. 1997, Bergin et al. 2000, Fairbairn and fi eld data limited in scale and extent. Until Dinsmore 2001, Naugle et al. 2001, Tewksbury recently, data suffi cient for analyzing regional et al. 2002), as well as scale-dependent and landscape-level habitat associations have responses to habitat characteristics (Pribil and not been available. Studies that published Picman 1997, Naugle et al. 1999). data on estuary-wide songbird distributions Bird relationships to patch size and shape (Hobson et al. 1986, Nur et al. 1997) did not have been studied in other habitats, especially generally contain corresponding information with regard to the process of habitat fragmenta- on critical habitat and landscape characteris- tion. Many researchers have evaluated island tics. Black Rail distribution patterns in the estu- biogeography (MacArthur and Wilson
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