Patrick D. Culbert

Land-Cover Change and Avian Diversity in the Conterminous United States

Changes in land use and land cover have affected and will continue to affect biological diversity worldwide. Yet, understanding the spatially extensive effects of land-cover change has been challenging because data that are consistent over space and time are lacking. We used the U.S. National Land Cover Dataset Land Cover Change Retrofit Product and North American Breeding Bird Survey data to examine land-cover change and its associations with diversity of birds with principally terrestrial life cycles (landbirds) in the conterminous United States. We used mixed-effects models and model selection to rank associations by ecoregion. Land cover in 3.22% of the area considered in our analyses changed from 1992 to 2001, and changes in species richness and abundance of birds were strongly associated with land-cover changes. Changes in species richness and abundance were primarily associated with changes in nondominant types of land cover, yet in many ecoregions different types of land cover were associated with species richness than were associated with abundance. Conversion of natural land cover to anthropogenic land cover was more strongly associated with changes in bird species richness and abundance than persistence of natural land cover in nearly all ecoregions and different covariates were most strongly associated with species richness than with abundance in 11 of 17 ecoregions. Loss of grassland and shrubland affected bird species richness and abundance in forested ecoregions. Loss of wetland was associated with bird abundance in forested ecoregions. Our findings highlight the value of understanding changes in nondominant land cover types and their association with bird diversity in the United States.

The Influence of Vertical and Horizontal Habitat Structure on Nationwide Patterns of Avian Biodiversity

ABSTRACT. With limited resources for habitat conservation, the accurate identification of high-value avian habitat is crucial. Habitat structure affects avian biodiversity but is difficult to quantify over broad extents. Our goal was to identify which measures of vertical and horizontal habitat structure are most strongly related to patterns of avian biodiversity across the conterminous United States and to determine whether new measures of vertical structure are complementary to existing, primarily horizontal, measures. For 2,546 North American Breeding Bird Survey routes across the conterminous United States, we calculated canopy height and biomass from the National Biomass and Carbon Dataset (NBCD) as measures of vertical habitat structure and used land-cover composition and configuration metrics from the 2001 National Land Cover Database (NLCD) as measures of horizontal habitat structure. Avian species richness was calculated for each route for all birds and three habitat guilds. Avian species richness was significantly related to measures derived from both the NBCD and NLCD. The combination of horizontal and vertical habitat structure measures was most powerful, yielding high R2 values for nationwide models of forest (0.70) and grassland (0.48) bird species richness. New measures of vertical structure proved complementary to measures of horizontal structure. These data allow the efficient quantification of habitat structure over broad scales, thus informing better land management and bird conservation.

Combined effects of heat waves and droughts on avian communities across the conterminous United States

Increasing surface temperatures and climatic variability associated with global climate change are expected to produce more frequent and intense heat waves and droughts in many parts of the world. Our goal was to elucidate the fundamental, but poorly understood, effects of these extreme weather events on avian communities across the conterminous United States. Specifically, we explored: (1) the effects of timing and duration of heat and drought events, (2) the effects of jointly occurring drought and heat waves relative to these events occurring in isolation, and (3) how effects vary among functional groups related to nest location and migratory habit, and among ecoregions with differing precipitation and temperature regimes. Using data from remote sensing, meteorological stations, and the North American Breeding Bird Survey, we used mixed effects models to quantify responses of overall and functional group abundance to heat waves and droughts (occurring alone or in concert) at two key periods in the annual cycle of birds: breeding and post-fledging. We also compared responses among species with different migratory and nesting characteristics, and among 17 ecoregions of the conterminous United States. We found large changes in avian abundances related to 100-year extreme weather events occurring in both breeding and post-fledging periods, but little support for an interaction among time periods. We also found that jointly-, rather than individually-occurring heat waves and droughts were both more common and more predictive of abundance changes. Declining abundance was the only significant response to post-fledging events, while responses to breeding period events were larger but could be positive or negative. Negative responses were especially frequent in the western U.S., and among ground-nesting birds and Neotropical migrants, with the largest single-season declines (36%) occurring among ground-nesting birds in the desert Southwest. These results indicate the importance of functional traits, timing, and geography in determining avian responses to weather extremes. Because dispersal to other regions appears to be an important avian response, it may be essential to maintain habitat refugia in a more climatically variable future.

The Impact of Phenological Variation on Texture Measures of Remotely Sensed Imagery

Measures of image texture derived from remotely sensed imagery have proven useful in many applications. However, when using multitemporal imagery or multiple images to cover a large study area, it is important to understand how image texture measures are affected by surface phenology. Our goal was to characterize the robustness to phenological variation of common firstand second-order texture measures of satellite imagery. Three North American study sites were chosen to represent different biomes. At each site, a suite of image textures were calculated for three to four dates across the growing season. Texture measures were compared among dates to quantify their stability, and the stability of measures was also compared between biomes. Interseasonal variability of texture measures was high overall (mean interseasonal coefficient of variation = 0.79), indicating that care must be taken when using measures of texture at different phenological stages. Certain texture measures, such as first-order mean and entropy, as well as second-order homogeneity, entropy, and dissimilarity, were more robust to phenological change than other measures.

Conservation of Forest Birds: Evidence of a Shifting Baseline in Community Structure

Background Quantifying changes in forest bird diversity is an essential task for developing effective conservation actions. When subtle changes in diversity accumulate over time, annual comparisons may offer an incomplete perspective of changes in diversity. In this case, progressive change, the comparison of changes in diversity from a baseline condition, may offer greater insight because changes in diversity are assessed over longer periods of times. Our objectives were to determine how forest bird diversity has changed over time and whether those changes were associated with forest disturbance. Methodology/Principal Findings We used North American Breeding Bird Survey data, a time series of Landsat images classified with respect to land cover change, and mixed-effects models to associate changes in forest bird community structure with forest disturbance, latitude, and longitude in the conterminous United States for the years 1985 to 2006. We document a significant divergence from the baseline structure for all birds of similar migratory habit and nest location, and all forest birds as a group from 1985 to 2006. Unexpectedly, decreases in progressive similarity resulted from small changes in richness (<1 species per route for the 22-year study period) and modest losses in abundance (−28.7–−10.2 individuals per route) that varied by migratory habit and nest location. Forest disturbance increased progressive similarity for Neotropical migrants, permanent residents, ground nesting, and cavity nesting species. We also documented highest progressive similarity in the eastern United States. Conclusions/Significance Contemporary forest bird community structure is changing rapidly over a relatively short period of time (e.g., ∼22 years). Forest disturbance and forest regeneration are primary factors associated with contemporary forest bird community structure, longitude and latitude are secondary factors, and forest loss is a tertiary factor. Importantly, these findings suggest some regions of the United States may already fall below the habitat amount threshold where fragmentation effects become important predictors of forest bird community structure.

Housing development erodes avian community structure in U.S. protected areas

Protected areas are a cornerstone for biodiversity conservation, but they also provide amenities that attract housing development on inholdings and adjacent private lands. We explored how this development affects biodiversity within and near protected areas among six ecological regions throughout the United States. We quantified the effect of housing density within, at the boundary, and outside protected areas, and natural land cover within protected areas, on the proportional abundance and proportional richness of three avian guilds within protected areas. We developed three guilds from the North American Breeding Bird Survey, which included Species of Greatest Conservation Need, land cover affiliates (e.g., forest breeders), and synanthropic species associated with urban environments. We gathered housing density data for the year 2000 from the U.S. Census Bureau, and centered the bird data on this year. We obtained land cover data from the 2001 National Land Cover Database, and we used single- and multiple-variable analyses to address our research question. In all regions, housing density within protected areas was positively associated with the proportional abundance or proportional richness of synanthropes, and negatively associated with the proportional abundance or proportional richness of Species of Greatest Conservation Need. These relationships were strongest in the eastern forested regions and the central grasslands, where more than 70% and 45%, respectively, of the variation in the proportional abundance of synanthropes and Species of Greatest Conservation Need were explained by housing within protected areas. Furthermore, in most regions, housing density outside protected areas was positively associated with the proportional abundance or proportional richness of synanthropes and negatively associated with the proportional abundance of land cover affiliates and Species of Greatest Conservation Need within protected areas. However, these effects were weaker than housing within protected areas. Natural land cover was high with little variability within protected areas, and consequently, was less influential than housing density within or outside protected areas explaining the proportional abundance or proportional richness of the avian guilds. Our results indicate that housing development within, at the boundary, and outside protected areas impacts avian community structure within protected areas throughout the United States.

Variability in energy influences avian distribution patterns across the USA

Habitat transformations and climate change are among the most important drivers of biodiversity loss. Understanding the factors responsible for the unequal distribution of species richness is a major challenge in ecology. Using data from the North American Breeding Bird Survey to measure species richness and a change metric extracted from the MODerate resolution Imaging Spectroradiometer (MODIS), we examined the influence of energy variability on the geographic distribution of avian richness across the conterminous U.S. and in the different ecoregions, while controlling for energy availability. The analysis compared three groups of birds: all species, Neotropical migrants, and permanent residents. We found that interannual variability in available energy explained more than half of the observed variation in bird richness in some ecoregions. In particular, energy variability is an important factor in explaining the patterns of overall bird richness and of permanent residents, in addition to energy availability. Our results showed a decrease in species richness with increasing energy variability and decreasing energy availability, suggesting that more species are found in more stable and more productive environments. However, not all ecoregions followed this pattern. The exceptions might reflect other biological factors and environmental conditions. With more ecoclimatic variability predicted for the future, this study provides insight into how energy variability influences the geographical patterns of species richness.

Long‐term avian community response to housing development at the boundary of US protected areas: effect size increases with time

Summary 1. Biodiversity conservation is a primary function of protected areas. However, protected areas also attract people, and therefore, land use has intensified at the boundaries of these lands globally. In the USA, since the 1970s, housing growth at the boundaries (<1 km) of protected areas has increased at a rate far higher than on more distant private lands. Here, we designed our analyses to address our central hypothesis that increasing housing density in and near protected areas will increasingly alter their avian communities. 2. We quantified the relationship between abundance and richness of protected-area avian species of greatest conservation need, land-cover affiliates (e.g. species associated with natural land cover such as forest breeders) and synanthropes (e.g. species associated with humans) with housing density on the boundary of protected areas and on more distant private lands from 1970 to 2010 in three ecoregions of the USA. We accomplished this using linear mixedmodel analyses, data from the US Census Bureau and 90 routes of the North American Breeding Bird Survey. 3. Housing density at the boundary of protected areas tended to be strongly negatively related with the abundance and richness of species of greatest conservation need and landcover affiliates (upwards of 88% of variance explained) and strongly positively related with synanthropes (upwards of 83% of variance explained). The effect size of these relationships increased in most cases from 1970 to 2010 and was greatest in the densely developed eastern forests. In the more sparsely populated West, we found similar, though weaker, associations. 4. Housing density on private lands more distant from protected areas had similar, but more muted negative effects. 5. Synthesis and applications. Our results illustrate that as housing density has increased along the boundary of protected areas, the conservation benefit of these lands has likely diminished. We urge conservation planners to prioritize the purchase of private-land inholdings in order to maximize the extent of unfragmented natural lands within protected areas. Further, we strongly recommend that land-use planners implement boundary management strategies to alter the pattern of human access to protected areas, cluster development to concentrate the footprint of rural housing, and establish conservation agreements through local land trusts to buffer protected areas from the effects of development along protected-area boundaries. To maximize the conservation benefit of protected areas, we suggest that housing development should be restricted within 1 km of their boundaries.