Evaluating vegetation effects on animal demographics: the role of plant phenology and sampling bias

Plant phenological processes produce temporal variation in the height and cover of vegetation. Key aspects of animal life cycles, such as reproduction, often coincide with the growing season and therefore may inherently covary with plant growth. When evaluating the influence of vegetation variables on demographic rates, the decision about when to measure vegetation relative to the timing of demographic events is important to avoid confounding between the demographic rate of interest and vegetation covariates. Such confounding could bias estimated effect sizes or produce results that are entirely spurious. We investigated how the timing of vegetation sampling affected the modeled relationship between vegetation structure and nest survival of greater sage-grouse (Centrocercus urophasianus), using both simulated and observational data. We used the height of live grasses surrounding nests as an explanatory covariate, and analyzed its effect on daily nest survival. We compared results between models that included grass height measured at the time of nest fate (hatch or failure) with models where grass height was measured on a standardized date - that of predicted hatch date. Parameters linking grass height to nest survival based on measurements at nest fate produced more competitive models, but slope coefficients of grass height effects were biased high relative to truth in simulated scenarios. In contrast, measurements taken at predicted hatch date accurately predicted the influence of grass height on nest survival. Observational data produced similar results. Our results demonstrate the importance of properly considering confounding between demographic traits and plant phenology. Not doing so can produce results that are plausible, but ultimately inaccurate.