Restored off-channel pond habitats create thermal regime diversity and refuges within a Mediterranean-climate watershed

Cool-water habitats provide increasingly vital refuges for cold-water fish living on the margins of their historical ranges; consequently, efforts to enhance or create cool-water habitat are becoming a major focus of river restoration practices. However, the effectiveness of restoration projects for providing thermal refuge and creating diverse temperature regimes at the watershed scale remains unclear. In the Klamath River in northern California, the Karuk Tribe Fisheries Program, the Mid-Klamath Watershed Council, and the U.S. Forest Service constructed a series of off-channel ponds that recreate floodplain habitat and support juvenile coho salmon (Oncorhynchus kisutch) and steelhead (O. mykiss) along the Klamath River and its tributaries. We instrumented these ponds and applied multivariate autoregressive time series models of fine-scale temperature data from ponds, tributaries, and the mainstem Klamath River to assess how off-channel ponds contributed to thermal regime diversity and thermal refuge habitat in the Klamath riverscape. Our analysis demonstrated that ponds provide diverse thermal habitats that are significantly cooler than creek or mainstem river habitats, even during severe drought. Wavelet analysis of long-term (10 years) temperature data indicated that thermal buffering (i.e. dampening of diel variation) increased over time but was disrupted by drought conditions in 2021. Our analysis demonstrates that in certain situations, human-made off-channel ponds can increase thermal diversity in modified riverscapes even during drought conditions, potentially benefiting floodplain-dependent cold-water species. Restoration actions that create and maintain thermal regime diversity and thermal refuges will become an essential tool to conserve biodiversity in climate-sensitive watersheds.