From Wiens to Robel: A Review of Grassland-Bird Habitat Selection

ABSTRACT Efforts to stabilize or increase grassland bird populations require identification of suitable habitat as a first step. Although the number of studies examining grassland-bird habitat selection has increased substantially in recent years, much uncertainty exists regarding local-scale habitat variables that researchers should consider. We reviewed 57 studies and identified important vegetation features correlated with grassland bird abundance, density, occurrence, and nest and territory selection. Our objectives were to 1) guide future studies of grassland-bird habitat use by providing a reduced set of relevant vegetation characteristics, 2) challenge researchers to critically think about what variables to consider, and 3) highlight the need to include consistent definitions of terms used to describe grassland bird habitat. We identified 9 variables that were important predictors of habitat use by grassland birds: coverage of bare ground (important in 50% of the instances where it was included), grass (34% of instances), dead vegetation (33% of instances), forbs (31% of instances), and litter (29% of instances), along with an index of vegetation density (39% of instances) and volume (39% of instances), litter depth (36% of instances), and vegetation height (41% of instances). Only 25% of studies provided information on effects sizes and measures of variance. Furthermore, definitions of measured habitat variables were not consistent among studies. We provide definitions of the 9 important variables and implore authors to report effect size and measures of variance. Standardization of terms and reporting of meaningful results will facilitate replication of wildlife research and enhance our ability to recognize general patterns that emerge from observational studies of habitat use.     

Habitat use and movement patterns by dependent and independent juvenile Grasshopper Sparrows during the post‐fledging period

The post‐fledging period is a critical life stage for young grassland birds. Habitat selection by recently fledged birds may differ from that of adults and may change as juveniles transition from the care and protection of parents to independence. To describe patterns of habitat selection during these important life stages, we studied habitat use by juvenile Grasshopper Sparrows (Ammodramus savannarum) in a Conservation Reserve Program grassland in Maryland. We used radio‐telemetry to track daily movement patterns of two age classes of Grasshopper Sparrows during the post‐fledging period. Sparrows were classified as either dependent (<32‐d‐old) or independent (≥32‐d‐old). We characterized the vegetation at 780 vegetation plots (390 plots where birds were located and 390 paired random plots). Microhabitats where dependent birds were found had significantly more bare ground, litter, and plant species richness than paired random plots. In addition, dependent birds were found in plots with less bare ground, more warm‐season grass cover, more total vegetation cover, and more forb cover than plots used by independent birds. Plots where independent birds were located also had significantly more bare ground than random plots. Dependent birds are less able to escape from predators because their flight feathers are not fully grown so they may benefit from remaining in areas of greater vegetation cover. However, juveniles transitioning from dependence to independence must forage on their own, possibly explaining their increased use of more open areas where foraging may be easier. To properly manage habitat for grassland birds, management strategies must consider the changing needs of birds during different stages of development. Our results highlight the importance of diverse grassland ecosystems for juvenile grassland birds during the transition to independence.     

Bird Communities and Biomass Yields in Potential Bioenergy Grasslands

Demand for bioenergy is increasing, but the ecological consequences of bioenergy crop production on working lands remain unresolved. Corn is currently a dominant bioenergy crop, but perennial grasslands could produce renewable bioenergy resources and enhance biodiversity. Grassland bird populations have declined in recent decades and may particularly benefit from perennial grasslands grown for bioenergy. We asked how breeding bird community assemblages, vegetation characteristics, and biomass yields varied among three types of potential bioenergy grassland fields (grass monocultures, grass-dominated fields, and forb-dominated fields), and assessed tradeoffs between grassland biomass production and bird habitat. We also compared the bird communities in grassland fields to nearby cornfields. Cornfields had few birds compared to perennial grassland fields. Ten bird Species of Greatest Conservation Need (SGCN) were observed in perennial grassland fields. Bird species richness and total bird density increased with forb cover and were greater in forb-dominated fields than grass monocultures. SGCN density declined with increasing vertical vegetation density, indicating that tall, dense grassland fields managed for maximum biomass yield would be of lesser value to imperiled grassland bird species. The proportion of grassland habitat within 1 km of study sites was positively associated with bird species richness and the density of total birds and SGCNs, suggesting that grassland bioenergy fields may be more beneficial for grassland birds if they are established near other grassland parcels. Predicted total bird density peaked below maximum biomass yields and predicted SGCN density was negatively related to biomass yields. Our results indicate that perennial grassland fields could produce bioenergy feedstocks while providing bird habitat. Bioenergy grasslands promote agricultural multifunctionality and conservation of biodiversity in working landscapes.     

Dung‐associated arthropods influence foraging ecology and habitat selection in Black‐necked Cranes (Grus nigricollis) on the Qinghai–Tibet Plateau

Variation in grassland vegetation structure influences the habitat selection of insectivorous birds. This variation presents a trade‐off for insectivorous predators: Arthropod abundance increases with vegetation height and heterogeneity, but access to arthropod prey items decreases. In contrast, grazing by large herbivores reduces and homogenizes vegetation, decreasing total arthropod abundance and diversity. However, the presence of livestock dung may help counteract the overall reduction in invertebrates by increasing arthropods associated with dung. It is unclear, however, how the presence of arthropod prey in dung contributes to overall habitat selection for insectivorous birds or how dung‐associated arthropods affect trade‐offs between vegetation structure, arthropod abundance, and access to prey. To explore these relationships, we studied habitat selection of the Black‐necked Crane (Grus nigricollis), a large omnivorous bird that breeds on the Qinghai–Tibet Plateau. We assessed the relationships between habitat selection of cranes and vegetation structure, arthropod abundance, and the presence of yak dung. We found that Black‐necked Cranes disproportionately foraged in grassland patches with short sward height, low sward height heterogeneity, and high numbers of dry yak dung, despite these habitats having lower total arthropod abundance. Although total arthropod abundance is lower, these habitats are supplemented with dry yak dung, which are associated with coleopteran larvae, making dung pats an indicator of food resources for breeding Black‐necked Cranes. Coleopteran adults and larvae in yak dung appear to be an important factor influencing the habitat selection of Black‐necked Cranes and should be considered when assessing grassland foraging trade‐offs of insectivorous birds. This research provides new insights into the role of livestock dung in defining foraging habitats and resources for insectivorous predators.     

Determinants of bird populations in an urban area

The effects of suburb age and distance from remnant native vegetation on the species richness and density of bird populations in Canberra were studied from June 1982 to May 1983. Mean total number of bird species increased with age of the suburbs < 12 years old. The density of birds also increased with suburb age. The number of open forest and woodland species increased with the age of the suburbs but the number of grassland and exotic species remained largely unchanged. These increases in bird species richness and density represented a response to changes in habitat conditions over time and were not a direct response to the suburb age per se. Such habitat changes reflected the growth of trees and, to a lesser extent, the increased cover of shrubs. The numbers of open forest, woodland and grassland species decreased with distance of suburban sites from native vegetation areas. The number of exotic species, however, remained unaffected. The density of individual birds of grassland and exotic species increased with distance of sites from remnant native vegetation.     

The nest predator community of grassland birds responds to agroecosystem habitat at multiple scales

Nest predation is the leading cause of reproductive failure for grassland birds of conservation concern. Understanding variation in nest predation rates is complicated by the diverse assemblage of species known to prey on nests. As part of a long‐term study of grassland bird ecology, we monitored populations of predators known to prey on grassland bird nests. We used information theoretic approach to examine the predator community's association with habitat at multiple scales, including local vegetation structure of grassland patches, spatial attributes of grassland patches (size and shape), and landscape composition surrounding grassland patches (land cover within 400 and 1600 m). Our results confirmed that nest predators respond to habitat at multiple scales and different predator species respond to habitat in different ways. The most informative habitat models we selected included variability in local vegetation (CV in the density of forbs), local patch (area and edge‐to‐interior ratio), and landscape within a 1600 m buffer around grasslands (percent of land covered by human structures and development). As a separate question, we asked if models that incorporated information from multiple scales simultaneously might improve the ability to explain variation in the predator community. Multi‐ scale models were not consistently superior to models derived from variables focused at a single spatial scale. Our results suggest that minimizing human development on and surrounding conservation land and the management of the vegetation structure on grassland fragments both may benefit grassland birds by decreasing the risk of nest predation.     

Can ecosystem services be integrated with conservation? A case study of breeding waders on grassland

We review published studies to show that changes in soil moisture levels have significant impacts on a range of wading bird species that use UK lowland grassland, including wet grassland, and obtain their food predominantly by probing the soil. We examine both the hydrological and the ecological literature and assess how management options could alter (1) ecosystem services (via water quality and flooding) and (2) habitat quality for wading birds. The combination of biodiversity goals with broader ecosystem services has been widely advocated and we suggest that appropriate management at multiple scales (e.g. small‐scale: ponds; large‐scale: integrated washlands) could potentially provide both ecosystem services and habitat for wading grassland birds. However, there is only a limited base of evidence on which to assess the potential linkage between these two areas, particularly for non‐wading bird species. Future work should be directed at identifying (1) how crop yield, ecosystem services and biodiversity relate to each other, (2) the extent of land needed to be managed to benefit these multiple purposes and bring about measurable gain (e.g. one or two ponds may make significant inroads in reducing run‐off and pollution but make little difference to wading birds) and (3) solutions to the challenges of setting up management options at large spatial scales (e.g. catchments).     

Airborne hyperspectral data predict Ellenberg indicator values for nutrient and moisture availability in dry grazed grasslands within a local agricultural landscape

Species-based ecological indices, such as Ellenberg indicators, reflect plant habitat preferences and can be used to describe local environment conditions. One disadvantage of using vegetation data as a substitute for environmental data is the fact that extensive floristic sampling can usually only be carried out at a plot scale within limited geographical areas. Remotely sensed data have the potential to provide information on fine-scale vegetation properties over large areas. In the present study, we examine whether airborne hyperspectral remote sensing can be used to predict Ellenberg nutrient (N) and moisture (M) values in plots in dry grazed grasslands within a local agricultural landscape in southern Sweden. We compare the prediction accuracy of three categories of model: (I) models based on predefined vegetation indices (VIs), (II) models based on waveband-selected VIs, and (III) models based on the full set of hyperspectral wavebands. We also identify the optimal combination of wavebands for the prediction of Ellenberg values. The floristic composition of 104 (4 m × 4 m grassland) plots on the Baltic island of Öland was surveyed in the field, and the vascular plant species recorded in the plots were assigned Ellenberg indicator values for N and M. A community-weighted mean value was calculated for N (mN) and M (mM) within each plot. Hyperspectral data were extracted from an 8 m × 8 m pixel window centred on each plot. The relationship between field-observed and predicted mean Ellenberg values was significant for all three categories of prediction models. The performance of the category II and III models was comparable, and they gave lower prediction errors and higher R² values than the category I models for both mN and mM. Visible and near-infrared wavebands were important for the prediction of both mN and mM, and shortwave infrared wavebands were also important for the prediction of mM. We conclude that airborne hyperspectral remote sensing can detect spectral differences in vegetation between grassland plots characterised by different mean Ellenberg N and M values, and that remote sensing technology can potentially be used to survey fine-scale variation in environmental conditions within a local agricultural landscape.     

Indices of bird-habitat preference from field surveys of birds and remote sensing of land cover: a study of south-eastern England with wider implications for conservation and biodiversity assessment

Aim This paper describes the development of novel indices of bird‐habitat preference to examine bird species’ use of habitats and their distributions relative to habitats. It assesses the implications for bird conservation regionally and the scope for biodiversity assessments generally. Location A 200 km by 400 km area of farmland with seminatural and urban areas, covering south‐eastern England. Methods Cluster analysis was used to link birds to landscapes. Cluster centroid coordinate values were processed to derive indices of bird‐habitat preference. Further developments assessed the relative values of individual habitats for birds. Results Clustering objectively linked birds to landscapes. Maps of the clusters showed strong regional patterns associated with distinctive habitat assemblages. Derived indices related bird species directly to individual habitats and habitats to birds. Even rare species and scarce habitats showed successful linkages, often to each other. Objective corroboration strongly supported the associations of coastal, wetland, urban and woodland birds and habitats; but, it suggested that farmland birds, whose numbers have nearly halved since 1977, may prefer alternative habitats. Main conclusions Land cover maps from remote sensing provide an effective way to link birds to habitats and vice versa. Thus, generalized habitat maps might be used to extrapolate localized or sample‐based bird observations or the results of autecological studies, helping to predict and understand bird distributions in the wider countryside. The weak links between farmland birds and farmland habitats in a region dominated by farming, suggests that reasons for the decline in farmland birds may be deep seated and thus hard to reverse. The procedures described are repeatable elsewhere and applicable more generally to evaluate landscapes and biodiversity. It is suggested that remote sensing could rarely be bettered as a means of assessing habitats, comprehensively, over wide areas, in most parts of the world.     

Disturbance-induced bird diversity in early successional habitats in the humid temperate region of northern Japan

The positive role of moderate natural disturbance is less known for a mobile organism such as birds, compared to sessile organisms. In the face of recent declines of grassland birds, it is necessary to identify the mechanism to maintain avian diversity in early successional open habitats in different regions. In the humid temperate region, the predominant habitat type is woody vegetation. Therefore, bird communities were studied along vegetational succession using a chronosequence method. TWINSPAN identified three distinctive habitat types (barren, grass-fen and shrubland) and four habitat guilds of birds (pioneer, grassland, ubiquitous and shrub) in the study sites. Path analysis determined the direct and indirect effects of disturbance on the habitat guilds of birds. Short and intermediate intervals of the water disturbance are important to pioneer guilds and grass-fen habitat, respectively. The frequent occurrence of large animals had a negative impact on grassland guild. It suggests that grassland birds less tolerate cattle grazing or human activities in a region with intensive land use. The chronosequence study revealed the dynamic nature of the bird community. Birds occupy habitats following the successional sere of vegetation, responding to animal occurrence and human activities in various manners.     

Relationships between vegetation and bird community composition in grasslands of the Serengeti

The Serengeti–Mara ecosystem holds one of the largest natural grasslands of the world, which is well known for its large herds of mammals. However, the bird community structure of these grasslands has hardly ever been studied. For the first time, a large‐scale study on grassland bird communities has been conducted in all typical open grasslands of the Serengeti National Park. We used ten grassland plots representing a gradient of increasing vegetation height and also including shrubs and trees to analyse the influence of vegetation structure on the composition of grassland bird communities. Three communities of breeding birds were identified in relation to (a) short grass, (b) intermediate and long grass and (c) wooded grasslands. The bird communities of intermediate, long and wooded grassland were very similar, because of identical dominant bird species. Our results suggest that breeding birds of East African grasslands exhibit two contrasting habitat relationships: (1) birds that are restricted to short grass, and (2) species that are tolerant to vegetational shifts from intermediate grass via long grass to early stages of woody vegetation. Because the vegetation is often driven beyond this range in managed tropical grasslands through high grazing pressure or through shrub encroachment, we expect that the long‐grass bird communities will not generally resist anthropogenic disturbance.     

Relationship between avian range limits and plant transition zones in Maine

Aim To determine if vegetation complexity associated with transition zones may be a contributing factor affecting bird species distributions in Maine, USA, and in increased numbers of bird species at about 45° north latitude in northeastern North America. Location Maine, USA; North America north of Mexico. Methods We delineated the ranges within Maine (86,156 km²) of 186 bird species and 240 woody plants using literature and expert review. Maps showing species richness and numbers of range limits, at 324 km² resolution, were developed for woody plants and groups of breeding birds: forest specialists, forest generalists, and those that used barren and urban habitats, early successional areas, and wetlands or open water. Two plant transition zones for Maine were identified previously, with the north–south transition zone mapped across eastern North America. Patterns in bird distribution maps were compared to woody plant maps and to transition zones. Results When the distributions of forest specialists were compared to the north–south vegetation transition zone in Maine, they were spatially coincident, but were not for other groups. Forest specialists had more species with range limits in the state (61%) than generalists (13%) or any other group. At a continental‐scale, the vegetation transition zone within eastern North America agreed fairly well with the areas of highest bird richness. Main conclusions A bird transition zone occurs in Maine and across eastern North America, akin to and overlapping the vegetation transition zone. Seasonality is likely the primary source of the inverse gradient in bird richness in the eastern USA, as reported by others. However, vegetation structure and habitat selection at very broad spatial scales appear to contribute to the reversed gradient. North of the vegetation transition zone, forest structure is simpler and coniferous forests more dominant, and this may contribute to reduced bird species richness. However, the northern (> 49°) typical gradient in bird species richness has been related to many hypotheses, and several are likely involved in the genesis of the gradient.     

Bird assemblages in fragmented agricultural landscapes: the role of small brigalow remnants and adjoining land uses

Agricultural intensification typically leads to changes in bird diversity and community composition, with fewer species and foraging guilds present in more intensively managed parts of the landscape. In this study, we compare bird communities in small (2–32 ha) brigalow (Acacia harpophylla) remnants with those in adjacent uncultivated grassland, previously cultivated grassland and current cropland, to determine the contribution of different land uses to bird diversity in the agricultural landscape. Twenty remnant brigalow patches and adjacent agricultural (‘matrix’) areas in southern inland Queensland, Australia were sampled for bird composition and habitat characteristics. The richness, abundance and diversity of birds were all significantly higher in brigalow remnants than in the adjacent matrix of cropping and grassland. Within the matrix, species richness and diversity were higher in uncultivated grasslands than in current cultivation or previously cultivated grasslands. Forty-four percent of bird species were recorded only in brigalow remnants and 78% of species were recorded in brigalow and at least one other land management category. Despite high levels of landscape fragmentation and modification, small patches of remnant brigalow vegetation provide important habitat for a unique and diverse assemblage of native birds. The less intensively managed components of the agricultural matrix also support diverse bird assemblages and thus, may be important for local and regional biodiversity conservation.     

Bird community response to filter strips in Maryland

Filter strips are strips of herbaceous vegetation planted along agricultural field margins adjacent to streams or wetlands and are designed to intercept sediment, nutrients, and agrichemicals. Roughly 16,000 ha of filter strips have been established in Maryland through the United States Department of Agriculture's Conservation Reserve Enhancement Program. Filter strips often represent the only uncultivated herbaceous areas on farmland in Maryland and therefore may be important habitat for early-successional bird species. Most filter strips in Maryland are planted to either native warm-season grasses or cool-season grasses and range in width from 10.7 m to 91.4 m. From 2004 to 2007 we studied the breeding and wintering bird communities in filter strips adjacent to wooded edges and non-buffered field edges and the effect that grass type and width of filter strips had on bird community composition. We used 5 bird community metrics (total bird density, species richness, scrub-shrub bird density, grassland bird density, and total avian conservation value), species-specific densities, nest densities, and nest survival estimates to assess the habitat value of filter strips for birds. Breeding and wintering bird community metrics were greater in filter strips than in non-buffered field edges but did not differ between cool-season and warm-season grass filter strips. Most breeding bird community metrics were negatively related to the percent cover of orchardgrass (Dactylis glomerata) in ≥1 yr. Breeding bird density was greater in narrow (<30 m) compared to wide (>60 m) filter strips. Our results suggest that narrow filter strips adjacent to wooded edges can provide habitat for many bird species but that wide filter strips provide better habitat for grassland birds, particularly obligate grassland species. If bird conservation is an objective, avoid planting orchardgrass in filter strips and reduce or eliminate orchardgrass from filter strips through management practices. © 2011 The Wildlife Society.     

Mapping migratory bird prevalence using remote sensing data fusion

Background

Improved maps of species distributions are important for effective management of wildlife under increasing anthropogenic pressures. Recent advances in lidar and radar remote sensing have shown considerable potential for mapping forest structure and habitat characteristics across landscapes. However, their relative efficacies and integrated use in habitat mapping remain largely unexplored. We evaluated the use of lidar, radar and multispectral remote sensing data in predicting multi-year bird detections or prevalence for 8 migratory songbird species in the unfragmented temperate deciduous forests of New Hampshire, USA.

Methodology and Principal Findings

A set of 104 predictor variables describing vegetation vertical structure and variability from lidar, phenology from multispectral data and backscatter properties from radar data were derived. We tested the accuracies of these variables in predicting prevalence using Random Forests regression models. All data sets showed more than 30% predictive power with radar models having the lowest and multi-sensor synergy (“fusion”) models having highest accuracies. Fusion explained between 54% and 75% variance in prevalence for all the birds considered. Stem density from discrete return lidar and phenology from multispectral data were among the best predictors. Further analysis revealed different relationships between the remote sensing metrics and bird prevalence. Spatial maps of prevalence were consistent with known habitat preferences for the bird species.

Conclusion and Significance

Our results highlight the potential of integrating multiple remote sensing data sets using machine-learning methods to improve habitat mapping. Multi-dimensional habitat structure maps such as those generated from this study can significantly advance forest management and ecological research by facilitating fine-scale studies at both stand and landscape level.     

Influence of habitat modification by livestock on páramo bird abundance in southern Andes of Ecuador

Livestock grazing on natural grasslands is widespread with negative consequences to biodiversity. In the High Andes, páramo grassland is a distinctive ecosystem where the influence of livestock grazing on páramo birds is poorly documented. We assessed the influence of habitat modification of páramo grassland related to livestock grazing on bird habitat guilds in the southern Andes of Ecuador. We recorded birds occurring along transects located in areas which showed a gradient (low to high) of grazing pressure. We found a decrease in abundance of páramo specialists in transects with more grazing pressure. We interpret this habitat modification as loss of key habitat necessary for habitat-specialized birds.     

The effects of grassland management using fire on habitat occupancy and conservation of birds in a mosaic landscape

Prescribed burning is routinely used to improve grazing in Pyrenean rangelands affected by an overall trend of land abandonment. This study considers the environmental variables influencing habitat occupancy by birds and the consequences of the use of fire in range management for bird conservation. Bird use and habitat structure of 11 cover types, the result of specific management regimes, were monitored for two breeding seasons in a mosaic landscape. Three main gradients of avian composition, corresponding to tree cover, shrub volume and grazing intensity, were identified from canonical correspondence analysis. The structure of the bird community seemed more intensely affected by species-specific selection of cover types than by the birds' use of multiple patches. Out of a total of 10 bird species analysed by a simultaneous confidence intervals procedure, four species with an unfavourable conservation status in Europe (Emberiza cia, Lullula arborea, Saxicola torquata and Lanius collurio) preferred managed grassland. Three types of grassland with shrubs (derived from single or repeated burning) had the highest bird conservation index (taking into account specific status and abundance of the bird assemblage), whereas forests showed middle or low values. The relation (P = 0.054) of this index to the logarithm of the pastoral value (which includes density and grazing quality of grasses) in currently managed cover types suggests that the objectives of grassland recovery by appropriate management practices and those of bird conservation coincide in our study 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 sandwichensis), 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.