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Bayesian cross-validation for model evaluation and selection, with application to the North American Breeding Bird Survey

The analysis of ecological data has changed in two important ways over the last 15 years. The development and easy availability of Bayesian computational methods has allowed and encouraged the fitting of complex hierarchical models. At the same time, there has been increasing emphasis on acknowledging and accounting for model uncertainty. Unfortunately, the ability to fit complex models has outstripped the development of tools for model selection and model evaluation: familiar model selection tools such as Akaike's information criterion and the deviance information criterion are widely known to be inadequate for hierarchical models. In addition, little attention has been paid to the evaluation of model adequacy in context of hierarchical modeling, i.e., to the evaluation of fit for a single model. In this paper, we describe Bayesian cross-validation, which provides tools for model selection and evaluation. We describe the Bayesian predictive information criterion and a Bayesian approximation to the BPIC known as the Watanabe-Akaike information criterion. We illustrate the use of these tools for model selection, and the use of Bayesian cross-validation as a tool for model evaluation, using three large data sets from the North American Breeding Bird Survey.

Ecology

Population trends for North American winter birds based on hierarchical models

Managing widespread and persistent threats to birds requires knowledge of population dynamics at large spatial and temporal scales. For over 100 yrs, the Audubon Christmas Bird Count (CBC) has enlisted volunteers in bird monitoring efforts that span the Americas, especially southern Canada and the United States. We employed a Bayesian hierarchical model to control for variation in survey effort among CBC circles and, using CBC data from 1966 to 2013, generated early-winter population trend estimates for 551 species of birds. Selecting a subset of species that do not frequent bird feeders and have ≥25% range overlap with the distribution of CBC circles (228 species) we further estimated aggregate (i.e., across species) trends for the entire study region and at the level of states/provinces, Bird Conservation Regions, and Landscape Conservation Cooperatives. Moreover, we examined the relationship between ten biological traits—range size, population size, migratory strategy, habitat affiliation, body size, diet, number of eggs per clutch, age at sexual maturity, lifespan, and tolerance of urban/suburban settings—and CBC trend estimates. Our results indicate that 68% of the 551 species had increasing trends within the study area over the interval 1966–2013. When trends were examined across the subset of 228 species, the median population trend for the group was 0.9% per year at the continental level. At the regional level, aggregate trends were positive in all but a few areas. Negative population trends were evident in lower latitudes, whereas the largest increases were at higher latitudes, a pattern consistent with range shifts due to climate change. Nine of 10 biological traits were significantly associated with median population trend; however, none of the traits explained >34% of the deviance in the data, reflecting the indirect relationships between population trend estimates and species traits. Trend estimates based on the CBC are broadly congruent with estimates based on the North American Breeding Bird Survey, another large-scale monitoring program. Both of these efforts, conducted by citizen scientists, will be required going forward to ensure robust inference about population dynamics in the face of climate and land cover changes.

Ecosphere

Access to bird population data

Access to bird population data is critical for effective conservation planning and implementation. Although a tremendous volume of baseline data exists, it is often diffusely distributed and inaccessible to the resource manager and decision maker. A mechanism that facilitates assembly, documentation and delivery of avian data in a user-friendly manner is needed in order to integrate bird-related information resources across agencies and organizations. To address this fundamental need, the National Biological Information Infrastructure (NBII), in partnership with the U.S. Geological Survey's Patuxent Wildlife Research Center and the U.S. Fish and Wildlife Service, is developing a web-based interactive system that will focus on access to bird population and habitat data used in bird management and conservation. This system, known as the NBII Bird Conservation Node, will support planning and evaluation of bird conservation activities within the context of the North American Bird Conservation Initiative (NABCI), a framework for collaboration among organizations interested in bird conservation across North America. Initial development of the NBII Bird Conservation Node will focus on creating a prototype mapping application that will provide interactive access to data from the North American Breeding Bird Survey, the Colonial Waterbird Survey, the Breeding Waterfowl Population and Habitat Survey, and the Atlantic Flyway Mid-winter Waterfowl Survey. This prototype mapping application, to be available on-line at http://www.nbii.gov by Sep 2001, will lay the foundation for establishment of a Migratory Bird Data Center at Patuxent Wildlife Research Center, and will provide an opportunity for linking to and establishing partnerships with other sources of bird population and habitat data available over the Internet.

Book chapter

Atlantic Flyway review: Region V

Banding activities in Region V in 1975 were similar to those of 1974. The stations at Woodend and St. Michaels were discontinued, but a new station 5 miles south of St. Michaels was opened at Bellevue by Henry Armistead. For the locations of the various banding stations in Region V see the map in the 1974 report (North American Bird Bander 1:75-82). As in the past several years, the great majority of the birds banded were caught at the three full-time stations: Adventure in the Potomac Valley near the fall line, Damsite on the east shore of Chesapeake Bay, and Kiptopeke at the mouth of Chesapeake Bay. At these three stations, 17,244 birds were banded during the 1975 fall season; this is 77% of the regional total of 22,315 birds.

North American Bird Bander

Combined analysis of roadside and off-road breeding bird survey data to assess population change in Alaska

Management interest in North American birds has increasingly focused on species that breed in Alaska, USA, and Canada, where habitats are changing rapidly in response to climatic and anthropogenic factors. We used a series of hierarchical models to estimate rates of population change in 2 forested Bird Conservation Regions (BCRs) in Alaska based on data from the roadside North American Breeding Bird Survey (BBS) and the Alaska Landbird Monitoring Survey, which samples off-road areas on public resource lands. We estimated long-term (1993–2015) population trends for 84 bird species from the BBS and short-term (2003–2015) trends for 31 species from both surveys. Among the 84 species with long-term estimates, 11 had positive trends and 17 had negative trends in 1 or both BCRs; negative trends were primarily found among aerial insectivores and wetland-associated species, confirming range-wide negative continental trends for many of these birds. Three species with negative trends in the contiguous United States and southern Canada had positive trends in Alaska, suggesting different population dynamics at the northern edges of their ranges. Regional population trends within Alaska differed for several species, particularly those represented by different subspecies in the 2 BCRs, which are separated by rugged, glaciated mountain ranges. Analysis of the roadside and off-road data in a joint hierarchical model with shared parameters resulted in improved precision of trend estimates and suggested a roadside-related difference in underlying population trends for several species, particularly within the Northwestern Interior Forest BCR. The combined analysis highlights the importance of considering population structure, physiographic barriers, and spatial heterogeneity in habitat change when assessing patterns of population change across a landscape as broad as Alaska. Combined analysis of roadside and off-road survey data in a hierarchical framework may be particularly useful for evaluating patterns of population change in relatively undeveloped regions with sparse roadside BBS coverage.

Condor

Differential changes in the onset of spring across US National Wildlife Refuges and North American migratory bird flyways

Warming temperatures associated with climate change can have indirect effects on migratory birds that rely on seasonally available food resources and habitats that vary across spatial and temporal scales. We used two heat-based indices of spring onset, the First Leaf Index (FLI) and the First Bloom Index (FBI), as proxies of habitat change for the period 1901 to 2012 at three spatial scales: the US National Wildlife Refuge System; the four major bird migratory flyways in North America; and the seasonal ranges (i.e., breeding and non-breeding grounds) of two migratory bird species, Blue-winged Warbler ( Vermivora cyanoptera ) and Whooping Crane ( Grus americana ). Our results show that relative to the historical range of variability, the onset of spring is now earlier in 76% of all wildlife refuges and extremely early (i.e., exceeding 95% of historical conditions) in 49% of refuges. In all flyways but the Pacific, the rate of spring advance is generally greater at higher latitudes than at lower latitudes. This differential rate of advance in spring onset is most pronounced in the Atlantic flyway, presumably because of a “warming hole” in the southeastern US. Both FLI and FBI have advanced markedly in the breeding ranges–but not the non-breeding ranges–of the two selected bird species, albeit with considerable intra-range variation. Differences among species in terms of migratory patterns and the location and extent of seasonal habitats, as well as shifts in habitat conditions over time, may complicate predictions of the vulnerability of migratory birds to climate change effects. This study provides insight into how differential shifts in the phenology of disparate but linked habitats could inform local- to landscape-scale management strategies for the conservation of migratory bird populations.

PLoS ONE

Incorporating location uncertainty improves inference with stop-level North American Breeding Bird Survey data

Ecological models should account for uncertainty to be most effective and useful. Yet, uncertainty from model covariates—unlike that from other sources, such as sampling error or process variability—is seldom explicitly incorporated. This can cause underestimates of uncertainty to cascade through model parameter estimates, predictions, and downstream uses. Burner et al. proposed a method for quantifying uncertainty in covariates and incorporating it into models using informative Bayesian priors. This method was applied to stop-level Breeding Bird Survey (BBS) analyses, where land cover uncertainty at each stop arises from substantial stop location uncertainty. A limited validation of model-estimated land cover, using stops with known locations, indicated the method’s potential effectiveness, but it was not rigorously evaluated. We conduct a robust simulation-based test, generating stop locations, extracting land cover, and simulating bird communities across 210 BBS routes in the upper Midwest. We compare 3 models: a “known” model with true land cover, a “naive” model assuming consistent 800-m stop spacing, and a “full” model using informative priors to estimate land cover. Species parameter estimates and predicted prevalence patterns across gradients in land cover from the full model approached those of the known model and were substantially closer to the true values used in simulations relative to those from the naive model. Naive model parameters were more biased relative to the other models, and credible intervals of predicted species prevalence rarely included the true simulated values. The full model also produced land cover covariate estimates closer to true simulation values relative to the mean informative priors. Our results show that, for the BBS, informative priors enable more accurate stop-level analyses despite location uncertainty. In contrast, naive models that ignore this uncertainty yield poor inferences. More broadly, we demonstrate empirically the utility of informative priors to account for covariate uncertainty in ecological models.

Michigan, Minnesota, Wisconson

First-time observer effects in the North American Breeding Bird Survey

Currently the operational analysis of Breeding Bird Survey (BBS) data by the National Biological Service accounts for observer differences in estimating the trend for each route, but within-observer differences are not modeled. We tested for the existence of a form of within-observer differences in skill level, namely a change in ability to count birds of a given species after an observer's first year on a given route. An increase in ability could positively bias the trend estimate. Removal of an observer's first year of observation on each route for the period 1966 to 1991 resulted in lower average unweighted trend estimates for 415 of 459 species (90%). These reductions were statistically significant for 213 species (46%). The average reduction in trend was 1.8% change per year (SD = 5.4%). In route-regression analysis, route data are weighted by a measure of precision. Removing first-year observer counts reduced the weighted trend estimate for 275 of 416 species (66%), but differences generally were small.

The Auk

Trends in American kestrel counts from the North American Breeding Bird Survey

A 15-year summary of the BBS data suggests that continental numbers of American Kestrels have increased during 1966-1979. According to analyses of physiographic and state/province strata, some areas are largely responsible for the increase in the U.S. and southern Canada and that only in Illinois and Arkansas have kestrels declined. BBS data are too few in Florida to detect trends concerning F. s. paulus. Surveying on special Raptor Routes, on which volunteers looked specifically for raptors while retracing their BBS route, did not significantly increase the kestrel counts, but did improve detection rates.

Book chapter

The role of observer bias in the North American Breeding Bird Survey

Ornithologists sampling breeding bird populations are subject to a number of biases in bird recognition and identification. Using Breeding Bird Survey data, these biases are examined qualitatively and quantitatively, and their effects on counts are evaluated. Differences in hearing ability and degree of expertise are the major observer biases considered. Other, more subtle influences are also discussed, including unfamiliar species, resolution, imagination, similar songs and attitude and condition of observers. In most cases, welltrained observers are comparable in ability and their differences contribute little beyond sampling error. However, just as hearing loss can affect results, so can an unprepared observer. These biases are important because they can reduce the credibility of any bird population sampling effort. Care is advised in choosing observers and in interpreting and using results when observers of variable competence are involved.

Book chapter

Progress report on the North American Breeding Bird Survey

The Breeding Bird Survey has been monitoring bird population changes in the U.S.A. and Canada since 1966. Each cooperator makes 50 3-minute stops at 0.8-km intervals along randomly selected roadside routes in 4 to 4 1/2 hours on one morning in June of each year. Data from the 1500 to 1600 routes are entered on magnetic tape, edited for accuracy, and then used for statistical analysis of population change at four geographic levels: physiographic region; state or province; region (eastern, central and western); and the continent. Examples of use of the data are: (1) detection of year-to-year changes in continental or rcgional populations; (2) use of population indices to show long-term population changes; (3) range expansion and contraction at the state level, including some of the reasons for those changes and documenting the associated population shifts within the original range; (4) mapping distribution and relative abundance on either a state, regional or continental scale; (5) mapping changes in abundance; and(6) comparing breeding with winter distribution. Future potential uses will include estimates of total continental populations and indices of environmental quality.

Acta Ornithologica