Jill A. Shaffer

Patch Size and Landscape Effects on Density and Nesting Success of Grassland Birds

Abstract Current management recommendations for grassland birds in North America emphasize providing large patches of grassland habitat within landscapes that have few forest or shrubland areas. These Bird Conservation Areas are being proposed under the assumption that large patches of habitat in treeless landscapes will maintain viable populations of grassland birds. This assumption requires that patch size and landscape features affect density and nesting success of grassland birds, and that these effects are consistent among years and regions and across focal species. However, these assumptions have not yet been validated for grassland birds, and the relative importance of local vegetation structure, patch size, and landscape composition on grassland bird populations is not well known. In addition, factors influencing grassland bird nesting success have been investigated mostly in small-scale and short-duration studies. To develop management guidelines for grassland birds, we tested the spatial and temporal repeatability of the influence of patch size and landscape composition on density and nesting success of 3 grassland passerines, after controlling for local-scale vegetation structure, climate, and—when analyzing nest success—bird density. We conducted our study during 4 years (1998–2001) in 44 study plots that were set up in 3 regions of the northern tallgrass prairie in Minnesota and North Dakota, USA. In these study plots we measured density and nesting success of clay-colored sparrows (Spizella pallida), Savannah sparrows (Passerculus sandwichensis), and bobolinks (Dolichonyx oryzivorus). Statistical models indicated that density was influenced by patch size, landscape, region, and local vegetation structure more so than by local vegetation structure alone. Both magnitude and direction of the response of density to patch size varied among regions, years, and species. In contrast, the direction of landscape effects was consistent among regions, years, and between Savannah sparrows and bobolinks. In each species, this landscape effect was independent of patch size. Nesting success was not clearly influenced by patch size or landscape composition, and none of the factors that influenced avian density also influenced nesting success in any of the 3 species. General statements on “optimal habitat” for grassland birds should therefore be viewed cautiously. Instead, long-term studies in different regions as well as a deeper understanding of the local system are needed to determine which factors are most important for grassland birds in a particular area.

VARIABILITY IN VEGETATION EFFECTS ON DENSITY AND NESTING SUCCESS OF GRASSLAND BIRDS

Abstract The structure of vegetation in grassland systems, unlike that in forest systems, varies dramatically among years on the same sites, and among regions with similar vegetation. The role of this variation in vegetation structure on bird density and nesting success of grassland birds is poorly understood, primarily because few studies have included sufficiently large temporal and spatial scales to capture the variation in vegetation structure, bird density, or nesting success. To date, no large-scale study on grassland birds has been conducted to investigate whether grassland bird density and nesting success respond similarly to changes in vegetation structure. However, reliable management recommendations require investigations into the distribution and nesting success of grassland birds over larger temporal and spatial scales. In addition, studies need to examine whether bird density and nesting success respond similarly to changing environmental conditions. We investigated the effect of vegetation structure on the density and nesting success of 3 grassland-nesting birds: clay-colored sparrow (Spizella pallida), Savannah sparrow (Passerculus sand-wichensis), and bobolink (Dolichonyx oryzivorus) in 3 regions of the northern tallgrass prairie in 1998–2001. Few vegetation features influenced the densities of our study species, and each species responded differently to those vegetation variables. We could identify only 1 variable that clearly influenced nesting success of 1 species: clay-colored sparrow nesting success increased with increasing percentage of nest cover from the surrounding vegetation. Because responses of avian density and nesting success to vegetation measures varied among regions, years, and species, land managers at all times need to provide grasslands with different types of vegetation structure. Management guidelines developed from small-scale, short-term studies may lead to misrepresentations of the needs of grassland-nesting birds.

NESTING BIOLOGY OF THREE GRASSLAND PASSERINES IN THE NORTHERN TALLGRASS PRAIRIE

Abstract Basic nesting information on grassland passerines is needed for improving grassland bird management. Among the information needs are (1) the suitability of nesting habitat, (2) periods during the breeding season in which birds are most vulnerable to disturbances, and (3) how to fit grasslands into a prioritization scheme for conservation. Comparisons of nesting parameters among grassland species will help identify important management considerations. We describe and compare nest-site characteristics, nesting phenology, clutch size, hatching and fledging success, and brood parasitism by Brown-headed Cowbirds (Molothrus ater) for three grassland passerine species nesting in tallgrass prairie of northwestern Minnesota and southeastern North Dakota. During 1998–2002, we found 793 Clay-colored Sparrow (Spizella pallida), 687 Savannah Sparrow (Passerculus sandwichensis), and 315 Bobolink (Dolichonyx oryzivorus) nests. These species differed in many aspects of their breeding ecology. Clay-colored and Savannah sparrows initiated their nests almost 2 weeks earlier than Bobolinks, with peak nesting occurring in June. Clutch size was lower (3.77 ± 0.03 SE) for Clay-colored Sparrows than Savannah Sparrows (4.13 ± 0.05) and Bobolinks (5.25 ± 0.08). The number of host eggs hatched per nest was higher in Bobolinks (3.46 ± 0.20) than in Clay-colored Sparrows (2.52 ± 0.09) and Savannah Sparrows (2.41 ± 0.11), but the number of young fledged per Bobolink nest (1.97) was similar to that of Savannah Sparrows (2.01). Clay-colored Sparrows fledged only 1.35 host young per nest. Mayfield nest success was higher for Savannah Sparrows (31.4%) than for Clay-colored Sparrows (27.4%) or Bobolinks (20.7%). The main cause of nest failure was nest predation: predation in Clay-colored Sparrows (47.9%) was higher than in Savannah Sparrows (33.5%) but similar to Bobolinks (41.8%). Brood parasitism was lower in Clay-colored Sparrows (5.1%) than in Bobolinks (10.8%), and intermediate (6.7%) in Savannah Sparrows. Compared with most other studies, grassland bird nests in our study area were more successful and less frequently parasitized; thus, northwestern Minnesota and southeastern North Dakota appear to provide important breeding habitat for grassland birds.

Habitat and nesting of Le Conte's Sparrows in the northern tallgrass prairie

Abstract Little is known about the breeding biology of the Le Contes Sparrow (Ammodramus leconteii), probably because of its secretive nature. We provide new information on several aspects of Le Contes Sparrow breeding biology, including rates of nest parasitism by Brown-headed Cowbirds (Molothrus ater) and potential factors affecting breeding densities and nesting success of the species. Our study was conducted in the tallgrass prairie of northwestern Minnesota and southeastern North Dakota during 1998–2002. Breeding densities varied among years, but this variation was not clearly linked to climatic patterns. Vegetation had some influence on densities of Le Contes Sparrows; densities were highest in grasslands with moderate amounts of bare ground. Prairie patch size and the percentage of shrubs and trees in the landscape had no recognizable influence on density. Nesting success was highly variable among sites and years and increased slightly with distance from trees. Rates of nest parasitism were low (1 of 50 nests parasitized), and clutch sizes were similar to those of other studies of Le Contes Sparrows.

INFLUENCE OF TREES IN THE LANDSCAPE ON PARASITISM RATES OF GRASSLAND PASSERINE NESTS IN SOUTHEASTERN NORTH DAKOTA

Abstract. Woody vegetation has been linked to increased rates of Brown-headed Cowbird (Molothrus ater) parasitism for some grassland hosts. In northern North Dakota, however, studies reported that parasitism of grassland passerine nests was lower in landscapes with trees than in those without trees. We looked for evidence of this pattern elsewhere, using data from two studies conducted on the Sheyenne National Grassland in southeastern North Dakota. Specifically, we examined the probability of parasitism relative to percent tree cover within 2 km of a nest. We found a negative relationship for grassland passerine nests of all species tested. Our results support the suggestion that cowbirds are less likely to parasitize nests of grassland passerines where tree cover on the landscape is greater. This pattern could be explained by cowbirds switching to alternative hosts in woodlands, but this hypothesis needs further testing.

Effects of wind-energy facilities on breeding grassland bird distributions

The contribution of renewable energy to meet worldwide demand continues to grow. Wind energy is one of the fastest growing renewable sectors, but new wind facilities are often placed in prime wildlife habitat. Long-term studies that incorporate a rigorous statistical design to evaluate the effects of wind facilities on wildlife are rare. We conducted a before-after-control-impact (BACI) assessment to determine if wind facilities placed in native mixed-grass prairies displaced breeding grassland birds. During 2003-2012, we monitored changes in bird density in 3 study areas in North Dakota and South Dakota (U.S.A.). We examined whether displacement or attraction occurred 1 year after construction (immediate effect) and the average displacement or attraction 2-5 years after construction (delayed effect). We tested for these effects overall and within distance bands of 100, 200, 300, and >300 m from turbines. We observed displacement for 7 of 9 species. One species was unaffected by wind facilities and one species exhibited attraction. Displacement and attraction generally occurred within 100 m and often extended up to 300 m. In a few instances, displacement extended beyond 300 m. Displacement and attraction occurred 1 year after construction and persisted at least 5 years. Our research provides a framework for applying a BACI design to displacement studies and highlights the erroneous conclusions that can be made without the benefit of adopting such a design. More broadly, species-specific behaviors can be used to inform management decisions about turbine placement and the potential impact to individual species. Additionally, the avoidance distance metrics we estimated can facilitate future development of models evaluating impacts of wind facilities under differing land-use scenarios.

DOES BODY SIZE AFFECT A BIRD'S SENSITIVITY TO PATCH SIZE AND LANDSCAPE STRUCTURE?

Abstract ABSTRACT Larger birds are generally more strongly affected by habitat loss and fragmentation than are smaller ones because they require more resources and thus larger habitat patches. Consequently, conservation actions often favor the creation or protection of larger over smaller patches. However, in grassland systems the boundaries between a patch and the surrounding landscape, and thus the perceived size of a patch, can be indistinct. We investigated whether eight grassland bird species with different body sizes perceived variation in patch size and landscape structure in a consistent manner. Data were collected from surveys conducted in 44 patches of northern tallgrass prairie during 1998–2001. The response to patch size was very similar among species regardless of body size (density was little affected by patch size), except in the Greater Prairie-Chicken (Tympanuchus cupido), which showed a threshold effect and was not found in patches smaller than 140 ha. In landscapes containing 0%–30% woody vegetation, smaller species responded more negatively to increases in the percentage of woody vegetation than larger species, but above an apparent threshold of 30%, larger species were not detected. Further analyses revealed that the observed variation in responses to patch size and landscape structure among species was not solely due to body size per se, but to other differences among species. These results indicate that a stringent application of concepts requiring larger habitat patches for larger species appears to limit the number of grassland habitats that can be protected and may not always be the most effective conservation strategy.

Modeling effects of crop production, energy development and conservation-grassland loss on avian habitat

Birds are essential components of most ecosystems and provide many services valued by society. However, many populations have undergone striking declines as habitats have been lost or degraded by human activities. Terrestrial grasslands are vital habitat for birds in the North American Prairie Pothole Region (PPR), but grassland conversion and fragmentation from agriculture and energy-production activities have destroyed or degraded millions of hectares. Conservation grasslands can provide alternate habitat. In the United States, the Conservation Reserve Program (CRP) is the largest program maintaining conservation grasslands on agricultural lands, but conservation grasslands in the PPR have declined by over 1 million ha since the program’s zenith in 2007. We used an ecosystem-services model (InVEST) parameterized for the PPR to quantify grassland-bird habitat remaining in 2014 and to assess degradation status of this remaining habitat as influenced by crop and energy (i.e., oil, natural gas, and wind) production. We compared our resultant habitat-quality ratings to grassland-bird abundance data from the North American Breeding Bird Survey to confirm that ratings were related to grassland-bird abundance. Of the grassland-bird habitat remaining in 2014, about 18% was degraded by nearby crop production, whereas energy production degraded an additional 16%. We further quantified changes in availability of grassland-bird habitat under various land-cover scenarios representing incremental losses (10%, 25%, 50%, 75%, and 100%) of CRP grasslands from 2014 levels. Our model identified 1 million ha (9%) of remaining grassland-bird habitat in the PPR that would be lost or degraded if all CRP conservation grasslands were returned to crop production. In addition to direct losses, an economic climate favoring energy and commodity production over conservation has resulted in substantial degradation of remaining grassland-bird habitat across the PPR. Other grassland regions of the world face similar challenges in maintaining avian habitat.

Effects of wind-energy facilities on grassland bird distributions

The contribution of renewable energy to meet worldwide demand continues to grow. Wind energy is one of the fastest growing renewable sectors, but new wind facilities are often placed in prime wildlife habitat. Long-term studies that incorporate a rigorous statistical design to evaluate the effects of wind facilities on wildlife are rare. We conducted a before-after-control-impact (BACI) assessment to determine if wind facilities placed in native mixed-grass prairies displaced breeding grassland birds. During 2003-2012, we monitored changes in bird density in 3 study areas in North Dakota and South Dakota (U.S.A.). We examined whether displacement or attraction occurred 1 year after construction (immediate effect) and the average displacement or attraction 2-5 years after construction (delayed effect). We tested for these effects overall and within distance bands of 100, 200, 300, and >300 m from turbines. We observed displacement for 7 of 9 species. One species was unaffected by wind facilities and one species exhibited attraction. Displacement and attraction generally occurred within 100 m and often extended up to 300 m. In a few instances, displacement extended beyond 300 m. Displacement and attraction occurred 1 year after construction and persisted at least 5 years. Our research provides a framework for applying a BACI design to displacement studies and highlights the erroneous conclusions that can be made without the benefit of adopting such a design. More broadly, species-specific behaviors can be used to inform management decisions about turbine placement and the potential impact to individual species. Additionally, the avoidance distance metrics we estimated can facilitate future development of models evaluating impacts of wind facilities under differing land-use scenarios.

Effects of wind-energy facilities on breeding grassland bird distributions: Wind-Energy Effects on Grassland Birds

The contribution of renewable energy to meet worldwide demand continues to grow. Wind energy is one of the fastest growing renewable sectors, but new wind facilities are often placed in prime wildlife habitat. Long-term studies that incorporate a rigorous statistical design to evaluate the effects of wind facilities on wildlife are rare. We conducted a before-after-control-impact (BACI) assessment to determine if wind facilities placed in native mixed-grass prairies displaced breeding grassland birds. During 2003-2012, we monitored changes in bird density in 3 study areas in North Dakota and South Dakota (U.S.A.). We examined whether displacement or attraction occurred 1 year after construction (immediate effect) and the average displacement or attraction 2-5 years after construction (delayed effect). We tested for these effects overall and within distance bands of 100, 200, 300, and >300 m from turbines. We observed displacement for 7 of 9 species. One species was unaffected by wind facilities and one species exhibited attraction. Displacement and attraction generally occurred within 100 m and often extended up to 300 m. In a few instances, displacement extended beyond 300 m. Displacement and attraction occurred 1 year after construction and persisted at least 5 years. Our research provides a framework for applying a BACI design to displacement studies and highlights the erroneous conclusions that can be made without the benefit of adopting such a design. More broadly, species-specific behaviors can be used to inform management decisions about turbine placement and the potential impact to individual species. Additionally, the avoidance distance metrics we estimated can facilitate future development of models evaluating impacts of wind facilities under differing land-use scenarios.