Hierarchical modelling and estimation of abundance and population trends in metapopulation designs.

1. Population assessment in changing environments is challenging because factors governing abundance may also affect detectability and thus bias observed counts. We describe a hierarchical modelling framework for estimating abundance corrected for detectability in metapopulation designs, where observations of 'individuals' (e.g. territories) are replicated in space and time. We consider two classes of models; first, we regard the data as independent binomial counts and model abundance and detectability based on a product-binomial likelihood. Secondly, we use the more complex detection-non-detection data for each territory to form encounter history frequencies, and analyse the resulting multinomial/Poisson hierarchical model. Importantly, we extend both models to directly estimate population trends over multiple years. Our models correct for any time trends in detectability when assessing population trends in abundance. 2. We illustrate both models for a farmland and a woodland bird species, skylark Alauda arvensis and willow tit Parus montanus, by applying them to Swiss BBS data, where 268 1 km(2) quadrats were surveyed two to three times during 1999-2003. We fit binomial and multinomial mixture models where log(abundance) depended on year, elevation, forest cover and transect route length, and logit(detection) on year, season and search effort. 3. Parameter estimates were very similar between models with confidence intervals overlapping for most parameters. Trend estimates were similar for skylark (-0.074 +/- 0.041 vs. -0.047 +/- 0.019) and willow tit (0.044 +/- 0.046 vs. 0.047 +/- 0.018). As expected, the multinomial model gave more precise estimates, but also yielded lower abundance estimates for the skylark. This may be due to effects of territory misclassification (lumping error), which do not affect the binomial model. 4. Both models appear useful for estimating abundance and population trends free from distortions by detectability in metapopulation designs with temporally replicated observations. The ability to obtain estimates of abundance and population trends that are unbiased with respect to any time trends in detectability ought to be a strong motivation for the collection of replicate observation data.

[1]  A. Brix Bayesian Data Analysis, 2nd edn , 2005 .

[2]  David J. Spiegelhalter,et al.  WinBUGS user manual version 1.4 , 2003 .

[3]  Andrew Gelman,et al.  Data Analysis Using Regression and Multilevel/Hierarchical Models , 2006 .

[4]  C. Krebs Ecology: The Experimental Analysis of Distribution and Abundance , 1973 .

[5]  J. Andrew Royle N‐Mixture Models for Estimating Population Size from Spatially Replicated Counts , 2004, Biometrics.

[6]  S. T. Buckland,et al.  Estimating Animal Abundance , 2002 .

[7]  J. Andrew Royle,et al.  MODELING AVIAN ABUNDANCE FROM REPLICATED COUNTS USING BINOMIAL MIXTURE MODELS , 2005 .

[8]  J. Andrew Royle,et al.  ESTIMATING ABUNDANCE FROM REPEATED PRESENCE–ABSENCE DATA OR POINT COUNTS , 2003 .

[9]  J. Andrew Royle,et al.  HIERARCHICAL SPATIAL MODELS OF ABUNDANCE AND OCCURRENCE FROM IMPERFECT SURVEY DATA , 2007 .

[10]  K. Norris,et al.  Managing threatened species: the ecological toolbox, evolutionary theory and declining-population paradigm , 2004 .

[11]  W. Kendall,et al.  HOW SHOULD DETECTION PROBABILITY BE INCORPORATED INTO ESTIMATES OF RELATIVE ABUNDANCE , 2002 .

[12]  M. Conroy,et al.  Analysis and Management of Animal Populations , 2002 .

[13]  D. Noble,et al.  Developing indicators for European birds , 2005, Philosophical Transactions of the Royal Society B: Biological Sciences.

[14]  William A. Link,et al.  A Hierarchical Model for Regional Analysis of Population Change Using Christmas Bird Count Data, with Application to the American Black Duck , 2006 .

[15]  S. R. Searle,et al.  Generalized, Linear, and Mixed Models , 2005 .

[16]  David L. Borchers,et al.  Estimating Animal Abundance: Closed Populations , 2010 .

[17]  Marc Kéry,et al.  Monitoring programs need to take into account imperfect species detectability , 2004 .

[18]  S. Wood Generalized Additive Models: An Introduction with R , 2006 .

[19]  K. Pollock,et al.  EXPERIMENTAL ANALYSIS OF THE AUDITORY DETECTION PROCESS ON AVIAN POINT COUNTS , 2007 .

[20]  J. Andrew Royle,et al.  Inference About Species Richness and Community Structure Using Species-Specific Occupancy Models in the National Swiss Breeding Bird Survey MHB , 2009 .

[21]  Darryl I. MacKenzie,et al.  Occupancy as a surrogate for abundance estimation , 2004 .

[22]  Andrew Gelman,et al.  R2WinBUGS: A Package for Running WinBUGS from R , 2005 .

[23]  J. Andrew Royle,et al.  ESTIMATING SITE OCCUPANCY RATES WHEN DETECTION PROBABILITIES ARE LESS THAN ONE , 2002, Ecology.

[24]  M. Efford Density estimation in live‐trapping studies , 2004 .

[25]  J. Andrew Royle,et al.  Trend estimation in populations with imperfect detection , 2009 .

[26]  J. Andrew Royle,et al.  A Bayesian state-space formulation of dynamic occupancy models. , 2007, Ecology.

[27]  Kenneth H. Pollock,et al.  Bayesian spatial modeling of data from avian point count surveys , 2008 .

[28]  J Andrew Royle,et al.  A hierarchical model for spatial capture-recapture data. , 2008, Ecology.

[29]  R. Dorazio On the choice of statistical models for estimating occurrence and extinction from animal surveys. , 2007, Ecology.

[30]  W. Link,et al.  A HIERARCHICAL ANALYSIS OF POPULATION CHANGE WITH APPLICATION TO CERULEAN WARBLERS , 2002 .

[31]  Li Zhang,et al.  Modeling Unobserved Sources of Heterogeneity in Animal Abundance Using a Dirichlet Process Prior , 2008, Biometrics.

[32]  J. Andrew Royle,et al.  MODELING ABUNDANCE EFFECTS IN DISTANCE SAMPLING , 2004 .

[33]  Graeme Caughley,et al.  Directions in conservation biology , 1994 .

[34]  K. Pollock,et al.  A Field Evaluation of Distance Measurement Error in Auditory Avian Point Count Surveys , 2007 .

[35]  J. Andrew Royle,et al.  Modelling occurrence and abundance of species when detection is imperfect , 2005 .

[36]  J. Andrew Royle,et al.  Hierarchical Modeling and Inference in Ecology: The Analysis of Data from Populations, Metapopulations and Communities , 2008 .

[37]  M. Kéry Species Richness and Community Dynamics: A Conceptual Framework , 2011 .

[38]  Robert M Dorazio,et al.  Improving Removal‐Based Estimates of Abundance by Sampling a Population of Spatially Distinct Subpopulations , 2005, Biometrics.

[39]  M. Kéry,et al.  Estimating species richness: calibrating a large avian monitoring programme , 2005 .

[40]  D L Borchers,et al.  Spatially Explicit Maximum Likelihood Methods for Capture–Recapture Studies , 2008, Biometrics.

[41]  Marc Kéry,et al.  Estimating Abundance From Bird Counts: Binomial Mixture Models Uncover Complex Covariate Relationships , 2008 .

[42]  J. Nelder,et al.  Generalized Linear Models with Random Effects: Unified Analysis via H-likelihood , 2006 .

[43]  M. Karim Generalized Linear Models With Random Effects , 1991 .

[44]  Neil D. Burgess,et al.  Bird Census Techniques , 1992 .

[45]  Marc Kéry,et al.  Imperfect detection and its consequences for monitoring for conservation , 2008 .

[46]  J. Andrew Royle,et al.  Hierarchical models of animal abundance and occurrence , 2006 .

[47]  P. McCullagh,et al.  Generalized Linear Models , 1992 .