Accounting for individual-specific variation in habitat-selection studies: Efficient estimation of mixed-effects models using Bayesian or frequentist computation

Stefanie Muff, Johannes Signer, John Fieberg

Research output: Contribution to journalArticlepeer-review

76 Scopus citations


Popular frameworks for studying habitat selection include resource-selection functions (RSFs) and step-selection functions (SSFs), estimated using logistic and conditional logistic regression, respectively. Both frameworks compare environmental covariates associated with locations animals visit with environmental covariates at a set of locations assumed available to the animals. Conceptually, slopes that vary by individual, that is, random coefficient models, could be used to accommodate inter-individual heterogeneity with either approach. While fitting such models for RSFs is possible with standard software for generalized linear mixed-effects models (GLMMs), straightforward and efficient one-step procedures for fitting SSFs with random coefficients are currently lacking. To close this gap, we take advantage of the fact that the conditional logistic regression model (i.e. the SSF) is likelihood-equivalent to a Poisson model with stratum-specific fixed intercepts. By interpreting the intercepts as a random effect with a large (fixed) variance, inference for random-slope models becomes feasible with standard Bayesian techniques, or with frequentist methods that allow one to fix the variance of a random effect. We compare this approach to other commonly applied alternatives, including models without random slopes and mixed conditional regression models fit using a two-step algorithm. Using data from mountain goats (Oreamnos americanus) and Eurasian otters (Lutra lutra), we illustrate that our models lead to valid and feasible inference. In addition, we conduct a simulation study to compare different estimation approaches for SSFs and to demonstrate the importance of including individual-specific slopes when estimating individual- and population-level habitat-selection parameters. By providing coded examples using integrated nested Laplace approximations (INLA) and Template Model Builder (TMB) for Bayesian and frequentist analysis via the R packages R-INLA and glmmTMB, we hope to make efficient estimation of RSFs and SSFs with random effects accessible to anyone in the field. SSFs with individual-specific coefficients are particularly attractive since they can provide insights into movement and habitat-selection processes at fine-spatial and temporal scales, but these models had previously been very challenging to fit.

Original languageEnglish (US)
Pages (from-to)80-92
Number of pages13
JournalJournal of Animal Ecology
Issue number1
StatePublished - Jan 1 2020

Bibliographical note

Funding Information:
JF received partial support from the Minnesota Agricultural Experimental Station and the McKnight Foundation, and SM was supported by the Faculty of Science of the University of Zurich. The authors would like to thank Irene Weinberger for the permission to use the Eurasian otter dataset, and Benedikt Gehr for comments on an earlier version of the manuscript. We also thank three reviewers for their thoughtful input that greatly helped to improve this article.

Publisher Copyright:
© 2019 The Authors. Journal of Animal Ecology © 2019 British Ecological Society


  • conditional logistic regression
  • glmmTMB
  • integrated nested Laplace approximations (INLA)
  • multinomial regression
  • random effects
  • resource-selection functions
  • step-selection functions

PubMed: MeSH publication types

  • Journal Article
  • Research Support, Non-U.S. Gov't


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