Subalpine Meadow Flowering Phenology Responses to Climate Change: Integrating Experimental and Gradient Methods

We integrated experimental and natural gradient field methods to investigate effects of climate change and variability on flowering phenology of 11 subalpine meadow shrub, forb, and graminoid species in Gunnison County, Colorado (USA). At a subalpine meadow site, overhead electric radiant heaters advanced snowmelt date by 16 d and warmed and dried soil during the growing season. At three additional sites, a snow removal manipulation advanced snowmelt date by 7 d without altering growing season soil microclimate. We compared phenological responses to experimental climate change with responses to natural microclimate variability across spatial gradients at small and landscape scales, as well as across a temporal gradient from a separate study. Both manipulations significantly advanced timing of flowering for the group of species and for most species individually, closely paralleling responses of timing to natural spatial and temporal variability in snowmelt date. Snowmelt date singularly explained observed shifts in timing only in the earliest flowering species, Claytonia lanceolata. Among all other species except Artemisia tridentata var. vaseyana, the latest flowering species, a consistent combination of temperature-related microclimate factors (earlier snowmelt date, warmer soil temperatures, and decreased soil degree-days) substantially explained earlier timing. Both manipulations also extended flowering duration for the group of species, similar to species' responses to natural snowmelt variability at small spatial scales. However, only early flowering species displayed consistent, significant changes in duration, with extended duration related to earlier snowmelt or warmer spring soil temperatures. Soil moisture was generally not a significant explanatory factor for either timing or duration of flowering. Best-fit microclimate models explained an average of 82% of variation in timing but only 38% of variation in duration across species. Our research demonstrates the value of comparing and synthesizing results of multiple field methods within a single study. This integrated approach makes it easier to identify robust community-wide trends, as well as species-specific responses of phenology to climate change. The predicted short-term flowering phenology responses to temperature-related aspects of climate change may lead to longer term asynchronies in interspecific interactions, potentially altering population and evolutionary dynamics, community structure, and ecosystem functioning. Corresponding Editor: S. C. Pennings