Confronting Imperfect Detection: Behavior of Binomial Mixture Models under Varying Circumstances of Visits, Sampling Sites, Detectability, and Abundance, in Small-Sample Situations

Abstract Binomial mixture models (BMMs) have been increasingly applied to account for imperfect detection and to estimate abundance from count data, but their performance has not been thoroughly evaluated. Here, I conducted simulation experiments to examine parameter estimates in BMMs under various situations. I generated data by assuming that abundance followed a Poisson distribution with an expected value &lgr; and that the number of detected individuals followed a binomial distribution with an individual detection probability p. In simple simulations without covariates for &lgr; and p, when the number of sampling sites (n) was between 20 and 160, BMMs could recover &lgr; and p under the following conditions: 0.1≤&lgr;≤160 and p≥0.1. However, within these ranges of &lgr; and p, the estimates were variable under lower values of &lgr; and p, although the situation improved as n increased. When &lgr; and p are expected to exceed these ranges and the sample size is small, the results suggest that sampling and/or modeling designs should be reconsidered. I then conducted simulation experiments with covariates. I assumed that &lgr; increased with a covariate (x) across 20 sampling sites. I varied p, number of visits (v), and their dependency on a covariate. To compare BMMs with analyses that did not accommodate imperfect detection, I fitted ordinary Poisson generalized linear models to mean and maximum counts (GLMmean and GLMmax). The results showed that GLMmax was superior to GLMmean because GLMmean underestimated &lgr; when p was small. GLMmax underestimated a coefficient of the covariate (slope) when v was negatively correlated with x. BMMs successfully recovered true values of the intercepts, slopes, and &lgr; in most cases. However, when p and v were small, and when p and &lgr; were highly negatively correlated due to their inverse dependency on x, estimates from BMMs were more variable.

[1]  Ché Elkin,et al.  Modeling abundance using N-mixture models: the importance of considering ecological mechanisms. , 2009, Ecological applications : a publication of the Ecological Society of America.

[2]  Ç. Şekercioğlu,et al.  The Worldwide Variation in Avian Clutch Size across Species and Space , 2008, PLoS biology.

[3]  Geoffrey R. Geupel,et al.  Handbook of Field Methods for Monitoring Landbirds , 2012 .

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

[5]  Stephen T. Buckland,et al.  Estimating bird abundance: making methods work , 2008, Bird Conservation International.

[6]  Christopher K. Wikle,et al.  Hierarchical Bayesian Models for Predicting The Spread of Ecological Processes , 2003 .

[7]  Brett T McClintock,et al.  Unmodeled observation error induces bias when inferring patterns and dynamics of species occurrence via aural detections. , 2010, Ecology.

[8]  Jim Schieck,et al.  Biased detection of bird vocalizations affects comparisons of bird abundance among forested habitats , 1997 .

[9]  T. Saitoh,et al.  Harvest-based estimation of population size for Sika deer on Hokkaido Island, Japan , 2002 .

[10]  M. McCarthy,et al.  The influence of abundance on detectability , 2013 .

[11]  J. Andrew Royle,et al.  Modelling community dynamics based on species‐level abundance models from detection/nondetection data , 2011 .

[12]  Rick Bonney,et al.  The history of public participation in ecological research , 2012 .

[13]  Richard T. Reynolds,et al.  A Variable Circular-Plot Method for Estimating Bird Numbers , 1980 .

[14]  Katherine C. Kendall,et al.  Linking landscape characteristics to local grizzly bear abundance using multiple detection methods in a hierarchical model , 2011 .

[15]  H. Possingham,et al.  IMPROVING PRECISION AND REDUCING BIAS IN BIOLOGICAL SURVEYS: ESTIMATING FALSE‐NEGATIVE ERROR RATES , 2003 .

[16]  J. Lamarque,et al.  Global Biodiversity: Indicators of Recent Declines , 2010, Science.

[17]  Y. Yamaura,et al.  Effects of stand, landscape, and spatial variables on bird communities in larch plantations and deciduous forests in central Japan , 2008 .

[18]  L. Marini,et al.  Is the human population a large‐scale indicator of the species richness of ground beetles? , 2010 .

[19]  J. Nichols,et al.  Inferences About Landbird Abundance from Count Data: Recent Advances and Future Directions , 2009 .

[20]  Rick Bonney,et al.  The current state of citizen science as a tool for ecological research and public engagement , 2012 .

[21]  M. Betts,et al.  Point count summary statistics differentially predict reproductive activity in bird-habitat relationship studies , 2005, Journal of Ornithology.

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

[23]  David R. Anderson,et al.  LANDBIRD COUNTING TECHNIQUES: CURRENT PRACTICES AND AN ALTERNATIVE , 2002 .

[24]  Marc J. Mazerolle,et al.  LANDSCAPE CHARACTERISTICS INFLUENCE POND OCCUPANCY BY FROGS AFTER ACCOUNTING FOR DETECTABILITY , 2005 .

[25]  James D Nichols,et al.  Relaxing the closure assumption in occupancy models: staggered arrival and departure times. , 2013, Ecology.

[26]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[27]  J. Andrew Royle,et al.  Species richness and occupancy estimation in communities subject to temporary emigration. , 2009, Ecology.

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

[29]  K. Pollock,et al.  Factors affecting aural detections of songbirds. , 2007, Ecological applications : a publication of the Ecological Society of America.

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

[31]  J. Kerr,et al.  The Macroecological Contribution to Global Change Solutions , 2007, Science.

[32]  M. Kéry,et al.  Predicting species distributions from checklist data using site‐occupancy models , 2010 .

[33]  Paul Hendricks,et al.  A Fixed-Radius Point Count Method for Nonbreeding and Breeding-Season Use , 1986 .

[34]  J Andrew Royle,et al.  Generalized site occupancy models allowing for false positive and false negative errors. , 2006, Ecology.

[35]  J. Andrew Royle,et al.  Biodiversity of man-made open habitats in an underused country: a class of multispecies abundance models for count data , 2012, Biodiversity and Conservation.

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

[37]  J. Andrew Royle,et al.  Presence‐only modelling using MAXENT: when can we trust the inferences? , 2013 .

[38]  Y. Yamaura,et al.  Bird responses to broad-leaved forest patch area in a plantation landscape across seasons. , 2009 .

[39]  R. Swihart,et al.  Absent or undetected? Effects of non-detection of species occurrence on wildlife-habitat models , 2004 .

[40]  James S. Clark,et al.  Why environmental scientists are becoming Bayesians , 2004 .

[41]  Andrew Thomas,et al.  WinBUGS - A Bayesian modelling framework: Concepts, structure, and extensibility , 2000, Stat. Comput..

[42]  D. Lindenmayer,et al.  Fitting and Interpreting Occupancy Models , 2013, PloS one.

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

[44]  Douglas H. Johnson In Defense of Indices: The Case of Bird Surveys , 2008 .

[45]  F. Nakamura,et al.  A preliminary study on the effects of line and selective thinning on bird communities in Hokkaido, northern Japan , 2013, Journal of Forestry Research.

[46]  Kate E. Jones,et al.  What is macroecology? , 2012, Biology Letters.

[47]  Marc Kéry,et al.  Hierarchical modelling and estimation of abundance and population trends in metapopulation designs. , 2010, The Journal of animal ecology.

[48]  W. Jetz,et al.  The global diversity of birds in space and time , 2012, Nature.

[49]  Anne E. Magurran,et al.  Quantifying temporal change in biodiversity: challenges and opportunities , 2013, Proceedings of the Royal Society B: Biological Sciences.

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

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

[52]  Catherine A Calder,et al.  Accounting for uncertainty in ecological analysis: the strengths and limitations of hierarchical statistical modeling. , 2009, Ecological applications : a publication of the Ecological Society of America.

[53]  Marc Kéry,et al.  Towards the modelling of true species distributions , 2011 .

[54]  Jason M. Evans,et al.  Does accounting for imperfect detection improve species distribution models , 2011 .

[55]  Susanne A. Fritz,et al.  What ' s on the horizon for macroecology? , 2012 .

[56]  D. MacKenzie Occupancy Estimation and Modeling: Inferring Patterns and Dynamics of Species Occurrence , 2005 .

[57]  Matthew G Betts,et al.  Estimating thresholds in occupancy when species detection is imperfect. , 2011, Ecology.

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

[59]  Darryl I. MacKenzie,et al.  Designing occupancy studies: general advice and allocating survey effort , 2005 .

[60]  H. Higuchi,et al.  The effect of landscape contexts on wintering bird communities in rural Japan , 2005 .

[61]  Michael Schaub,et al.  Bayesian Population Analysis using WinBUGS: A Hierarchical Perspective , 2011 .

[62]  Richard B. Chandler,et al.  unmarked: An R Package for Fitting Hierarchical Models of Wildlife Occurrence and Abundance , 2011 .

[63]  Robert M. Dorazio,et al.  Occupancy estimation and the closure assumption , 2009 .

[64]  Len Thomas,et al.  Distance software: design and analysis of distance sampling surveys for estimating population size , 2009, The Journal of applied ecology.

[65]  S. Buckland Introduction to distance sampling : estimating abundance of biological populations , 2001 .

[66]  G. McKay,et al.  Encyclopedia of Animals , 2006 .