Abundance estimation in ecology is usually accomplished by capture-recapture, removal, or distance sampling methods. These may be hard to implement at large spatial scales. In contrast, binomial mixture models enable abundance estimation without individual identification, based simply on temporally and spatially replicated counts. Here, we evaluate mixture models using data from the national breeding bird monitoring program in Switzerland, where some 250 1-km 2 quadrats are surveyed using the territory mapping method three times during each breeding season. We chose eight species with contrasting distribution (wide-narrow), abundance (high-low), and detectability (easy-difficult). Abundance was modeled as a random effect with a Poisson or negative binomial distribution, with mean affected by forest cover, elevation, and route length. Detectability was a logitlinear function of survey date, survey date-by-elevation, and sampling effort (time per transect unit). Resulting covariate effects and parameter estimates were consistent with expectations. Detectability per territory (for three surveys) ranged from 0.66 to 0.94 (mean 0.84) for easy species, and from 0.16 to 0.83 (mean 0.53) for difficult species, depended on survey effort for two easy and all four difficult species, and changed seasonally for three easy and three difficult species. Abundance was positively related to route length in three high-abundance and one low-abundance (one easy and three difficult) species, and increased with forest cover in five forest species, decreased for two nonforest species, and was unaffected for a generalist species. Abundance estimates under the most parsimonious mixture models were between 1.1 and 8.9 (median 1.8) times greater than estimates based on territory mapping; hence, three surveys were insufficient to detect all territories for each species. We conclude that binomial mixture models are an important new approach for estimating abundance corrected for detectability when only repeated-count data are available. Future developments envisioned include estimation of trend, occupancy, and total regional abundance.
Correlative species distribution models are frequently used to predict species’ range shifts under climate change. However, climate variables often show high collinearity and most statistical approaches require the selection of one among strongly correlated variables. When causal relationships between species presence and climate parameters are unknown, variable selection is often arbitrary, or based on predictive performance under current conditions. While this should only marginally affect current range predictions, future distributions may vary considerably when climate parameters do not change in concert. We investigated this source of uncertainty using four highly correlated climate variables together with a constant set of landscape variables in order to predict current (2010) and future (2050) distributions of four mountain bird species in central Europe. Simulating different parameterization decisions, we generated a) four models including each of the climate variables singly, b) a model taking advantage of all variables simultaneously and c) an un‐weighted average of the predictions of a). We compared model accuracy under current conditions, predicted distributions under four scenarios of climate change, and – for one species – evaluated back‐projections using historical occurrence data. Although current and future variable‐correlations remained constant, and the models’ accuracy under contemporary conditions did not differ, future range predictions varied considerably in all climate change scenarios. Averaged models and models containing all climate variables simultaneously produced intermediate predictions; the latter, however, performed best in back‐projections. This pattern, consistent across different modelling methods, indicates a benefit from including multiple climate predictors in ambiguous situations. Variable selection proved to be an important source of uncertainty for future range predictions, difficult to control using contemporary information. Small, but diverging changes of climate variables, masked by constant overall correlation patterns, can cause substantial differences between future range predictions which need to be accounted for, particularly when outcomes are intended for conservation decisions.
Many estimation and inference problems arising from large‐scale animal surveys are focused on developing an understanding of patterns in abundance or occurrence of a species based on spatially referenced count data. One fundamental challenge, then, is that it is generally not feasible to completely enumerate (“census”) all individuals present in each sample unit. This observation bias may consist of several components, including spatial coverage bias (not all individuals in the population are exposed to sampling) and detection bias (exposed individuals may go undetected). Thus, observations are biased for the state variable (abundance, occupancy) that is the object of inference. Moreover, data are often sparse for most observation locations, requiring consideration of methods for spatially aggregating or otherwise combining sparse data among sample units. The development of methods that unify spatial statistical models with models accommodating non‐detection is necessary to resolve important spatial inference problems based on animal survey data. In this paper, we develop a novel hierarchical spatial model for estimation of abundance and occurrence from survey data wherein detection is imperfect. Our application is focused on spatial inference problems in the Swiss Survey of Common Breeding Birds. The observation model for the survey data is specified conditional on the unknown quadrat population size, N(s). We augment the observation model with a spatial process model for N(s), describing the spatial variation in abundance of the species. The model includes explicit sources of variation in habitat structure (forest, elevation) and latent variation in the form of a correlated spatial process. This provides a model‐based framework for combining the spatially referenced samples while at the same time yielding a unified treatment of estimation problems involving both abundance and occurrence. We provide a Bayesian framework for analysis and prediction based on the integrated likelihood, and we use the model to obtain estimates of abundance and occurrence maps for the European Jay (Garrulus glandarius), a widespread, elusive, forest bird. The naive national abundance estimate ignoring imperfect detection and incomplete quadrat coverage was 77 766 territories. Accounting for imperfect detection added approximately 18 000 territories, and adjusting for coverage bias added another 131 000 territories to yield a fully corrected estimate of the national total of about 227 000 territories. This is approximately three times as high as previous estimates that assume every territory is detected in each quadrat.
Species' assessments must frequently be derived from opportunistic observations made by volunteers (i.e., citizen scientists). Interpretation of the resulting data to estimate population trends is plagued with problems, including teasing apart genuine population trends from variations in observation effort. We devised a way to correct for annual variation in effort when estimating trends in occupancy (species distribution) from faunal or floral databases of opportunistic observations. First, for all surveyed sites, detection histories (i.e., strings of detection-nondetection records) are generated. Within-season replicate surveys provide information on the detectability of an occupied site. Detectability directly represents observation effort; hence, estimating detectability means correcting for observation effort. Second, site-occupancy models are applied directly to the detection-history data set (i.e., without aggregation by site and year) to estimate detectability and species distribution (occupancy, i.e., the true proportion of sites where a species occurs). Site-occupancy models also provide unbiased estimators of components of distributional change (i.e., colonization and extinction rates). We illustrate our method with data from a large citizen-science project in Switzerland in which field ornithologists record opportunistic observations. We analyzed data collected on four species: the widespread Kingfisher (Alcedo atthis) and Sparrowhawk (Accipiter nisus) and the scarce Rock Thrush (Monticola saxatilis) and Wallcreeper (Tichodroma muraria). Our method requires that all observed species are recorded. Detectability was <1 and varied over the years. Simulations suggested some robustness, but we advocate recording complete species lists (checklists), rather than recording individual records of single species. The representation of observation effort with its effect on detectability provides a solution to the problem of differences in effort encountered when extracting trend information from haphazard observations. We expect our method is widely applicable for global biodiversity monitoring and modeling of species distributions.
Mountain areas often hold special species communities, and they are high on the list of conservation concern. Global warming and changes in human land use, such as grazing pressure and afforestation, have been suggested to be major threats for biodiversity in the mountain areas, affecting species abundance and causing distribution shifts towards mountaintops. Population shifts towards poles and mountaintops have been documented in several areas, indicating that climate change is one of the key drivers of species’ distribution changes. Despite the high conservation concern, relatively little is known about the population trends of species in mountain areas due to low accessibility and difficult working conditions. Thanks to the recent improvement of bird monitoring schemes around Europe, we can here report a first account of population trends of 44 bird species from four major European mountain regions: Fennoscandia, UK upland, south‐western (Iberia) and south‐central mountains (Alps), covering 12 countries. Overall, the mountain bird species declined significantly (−7%) during 2002–2014, which is similar to the declining rate in common birds in Europe during the same period. Mountain specialists showed a significant −10% decline in population numbers. The slope for mountain generalists was also negative, but not significantly so. The slopes of specialists and generalists did not differ from each other. Fennoscandian and Iberian populations were on average declining, while in United Kingdom and Alps, trends were nonsignificant. Temperature change or migratory behaviour was not significantly associated with regional population trends of species. Alpine habitats are highly vulnerable to climate change, and this is certainly one of the main drivers of mountain bird population trends. However, observed declines can also be partly linked with local land use practices. More efforts should be undertaken to identify the causes of decline and to increase conservation efforts for these populations.
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