An efficient protocol for the global sensitivity analysis of stochastic ecological models

Stochastic simulation models requiring many input parameters are widely used to inform the management of ecological systems. The interpretation of complex models is aided by global sensitivity analysis, using simulations for distinct parameter sets sampled from multidimensional space. Ecologists typically analyze such output using an "emulator"; that is, a statistical model used to approximate the relationship between parameter inputs and simulation outputs and to derive sensitivity measures. However, it is typical for ad hoc decisions to be made regarding: (1) trading off the number of parameter samples against the number of simulation iterations run per sample, (2) determining whether parameter sampling is sufficient, and (3) selecting an appropriate emulator. To evaluate these choices, we coupled different sensitivity-analysis designs and emulators for a stochastic, 20-parameter model that simulated the re-introduction of a threatened species subject to predation and disease, and then validated the emulators against new output generated from the simulation model. Our results lead to the following sensitivity analysis-protocol for stochastic ecological models. (1) Run a single simulation iteration per parameter sample generated, even if the focal response is a probabilistic outcome, while sampling extensively across the parameter space. In contrast to designs that invested in many model iterations (tens to thousands) per parameter sample, this approach allowed emulators to capture the input-output relationship of the simulation model more accurately and also to produce sensitivity measures that were robust to variation inherent in the parameter-sampling stage. (2) Confirm that parameter sampling is sufficient, by emulating subsamples of the sensitivity-analysis output. As the subsample size is increased, the cross-validatory performance of the emulator and sensitivity measures derived from it should exhibit asymptotic behavior. This approach can also be used to compare candidate emulators and select an appropriate interaction depth. (3) If required, conduct further simulations for additional parameter samples, and then report sensitivity measures and illustrate key response curves using the selected emulator. This protocol will generate robust sensitivity measures and facilitate the interpretation of complex ecological models, while minimizing simulation effort.

[1]  Damien A. Fordham,et al.  Novel coupling of individual-based epidemiological and demographic models predicts realistic dynamics of tuberculosis in alien buffalo , 2012 .

[2]  H. Possingham,et al.  DO HARVEST REFUGES BUFFER KANGAROOS AGAINST EVOLUTIONARY RESPONSES TO SELECTIVE HARVESTING , 2004 .

[3]  Scott Ferson,et al.  Sensitivity analysis for models of population viability , 1995 .

[4]  M. Palmer,et al.  Quantitative Bioscience for the 21st Century , 2005 .

[5]  Barry W. Brook,et al.  Demographic sensitivity and persistence of the threatened white- and orange-bellied frogs of Western Australia , 2003, Population Ecology.

[6]  J Elith,et al.  A working guide to boosted regression trees. , 2008, The Journal of animal ecology.

[7]  G. De’ath The multinomial diversity model: linking Shannon diversity to multiple predictors. , 2012, Ecology.

[8]  R. Tibshirani,et al.  Generalized Additive Models , 1991 .

[9]  Thomas A A Prowse,et al.  An ecological regime shift resulting from disrupted predator-prey interactions in Holocene Australia. , 2014, Ecology.

[10]  J. Sabo,et al.  Population viability and species interactions : Life outside the single-species vacuum , 2008 .

[11]  T. Blackburn,et al.  A population model for predicting the successful establishment of introduced bird species , 2014, Oecologia.

[12]  Andrea Castelletti,et al.  Emulation techniques for the reduction and sensitivity analysis of complex environmental models , 2012, Environ. Model. Softw..

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

[14]  A. Saltelli,et al.  The role of sensitivity analysis in ecological modelling , 2007 .

[15]  Geoffrey R Hosack,et al.  Assessing model structure uncertainty through an analysis of system feedback and Bayesian networks. , 2008, Ecological applications : a publication of the Ecological Society of America.

[16]  Robert C. Lacy,et al.  Metamodels for Transdisciplinary Analysis of Wildlife Population Dynamics , 2013, PloS one.

[17]  I. Sobola,et al.  Global sensitivity indices for nonlinear mathematical models and their Monte Carlo estimates , 2001 .

[18]  Lynn A. Maguire,et al.  Sample Sizes for Minimum Viable Population Estimation , 1987 .

[19]  D. Lindenmayer,et al.  Modelling the viability of metapopulations of the endangered Leadbeater's possum in south-eastern Australia , 1995, Biodiversity & Conservation.

[20]  Hannu Toivonen,et al.  BAYESIAN ANALYSIS OF METAPOPULATION DATA , 2002 .

[21]  M. D. McKay,et al.  A comparison of three methods for selecting values of input variables in the analysis of output from a computer code , 2000 .

[22]  F. Ayala,et al.  Complexity in Ecology and Conservation: Mathematical, Statistical, and Computational Challenges , 2005 .

[23]  Pierre Petitgas,et al.  Combining sensitivity and uncertainty analysis to evaluate the impact of management measures with ISIS–Fish: marine protected areas for the Bay of Biscay anchovy (Engraulis encrasicolus) fishery , 2010 .

[24]  Christopher N. Johnson,et al.  Ecological and economic benefits to cattle rangelands of restoring an apex predator , 2015 .

[25]  H. Resit Akçakaya,et al.  Predictive accuracy of population viability analysis in conservation biology , 2000, Nature.

[26]  Sean M. McMahon,et al.  On using integral projection models to generate demographically driven predictions of species' distributions: development and validation using sparse data , 2014 .

[27]  Denis Valle,et al.  The importance of multimodel projections to assess uncertainty in projections from simulation models. , 2009, Ecological applications : a publication of the Ecological Society of America.

[28]  Stefan Finsterle,et al.  Making sense of global sensitivity analyses , 2014, Comput. Geosci..

[29]  S. Wood Fast stable restricted maximum likelihood and marginal likelihood estimation of semiparametric generalized linear models , 2011 .

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

[31]  C. Bradshaw,et al.  Minimum viable population size: A meta-analysis of 30 years of published estimates , 2007 .

[32]  Hiroyuki Yokomizo,et al.  Meta-models as a straightforward approach to the sensitivity analysis of complex models , 2013, Population Ecology.

[33]  Robert C. Lacy,et al.  VORTEX: a computer simulation model for population viability analysis , 1993 .

[34]  Christopher N. Johnson,et al.  No need for disease: testing extinction hypotheses for the thylacine using multi-species metamodels. , 2013, The Journal of animal ecology.

[35]  Lev R Ginzburg,et al.  Rules of thumb for judging ecological theories. , 2004, Trends in ecology & evolution.

[36]  Martin Drechsler,et al.  Sensitivity analysis of complex models , 1998 .

[37]  M. Shaffer Minimum Population Sizes for Species Conservation , 1981 .

[38]  Runze Li,et al.  Design and Modeling for Computer Experiments , 2005 .

[39]  Brett A. Melbourne,et al.  Extinction risk depends strongly on factors contributing to stochasticity , 2008, Nature.

[40]  Jon C. Helton,et al.  Implementation and evaluation of nonparametric regression procedures for sensitivity analysis of computationally demanding models , 2009, Reliab. Eng. Syst. Saf..

[41]  F. Vinatier,et al.  Explaining host–parasitoid interactions at the landscape scale: a new approach for calibration and sensitivity analysis of complex spatio-temporal models , 2012, Landscape Ecology.

[42]  Peter Arcese,et al.  Sensitivity Analyses of Spatial Population Viability Analysis Models for Species at Risk and Habitat Conservation Planning , 2009, Conservation biology : the journal of the Society for Conservation Biology.

[43]  Barry W. Brook,et al.  Pessimistic and Optimistic Bias in Population Viability Analysis , 2000 .

[44]  R. Levins The strategy of model building in population biology , 1966 .

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

[46]  M. Boyce Population Viability Analysis , 1992 .

[47]  Guillaume Marie,et al.  Extending the use of ecological models without sacrificing details: a generic and parsimonious meta‐modelling approach , 2014 .