Understanding recent biogeographic responses to climate change is fundamental for improving our predictions of likely future responses and guiding conservation planning at both local and global scales. Studies of observed biogeographic responses to 20th century climate change have principally examined effects related to ubiquitous increases in temperature – collectively termed a warming fingerprint. Although the importance of changes in other aspects of climate – particularly precipitation and water availability – is widely acknowledged from a theoretical standpoint and supported by paleontological evidence, we lack a practical understanding of how these changes interact with temperature to drive biogeographic responses. Further complicating matters, differences in life history and ecological attributes may lead species to respond differently to the same changes in climate. Here, we examine whether recent biogeographic patterns across California are consistent with a warming fingerprint. We describe how various components of climate have changed regionally in California during the 20th century and review empirical evidence of biogeographic responses to these changes, particularly elevational range shifts. Many responses to climate change do not appear to be consistent with a warming fingerprint, with downslope shifts in elevation being as common as upslope shifts across a number of taxa and many demographic and community responses being inconsistent with upslope shifts. We identify a number of potential direct and indirect mechanisms for these responses, including the influence of aspects of climate change other than temperature (e.g., the shifting seasonal balance of energy and water availability), differences in each taxon's sensitivity to climate change, trophic interactions, and land-use change. Finally, we highlight the need to move beyond a warming fingerprint in studies of biogeographic responses by considering a more multifaceted view of climate, emphasizing local-scale effects, and including a priori knowledge of relevant natural history for the taxa and regions under study.

Heterogeneous terrain in montane systems results in a decoupling of climatic gradients. Population dynamics across species' ranges in these heterogeneous landscapes are shaped by relationships between demographic rates and these interwoven climate gradients. Linking demography and climate variables across species' ranges refines our understanding of the underlying mechanisms of species' current and future ranges.We explored the importance of multiple microclimatic gradients in shaping individual demographic rates and population growth rates in 16 populations across the elevational distribution of an alpine plant (Ivesia lycopodioides var. scandularis). Using integral projection modeling, we ask how each rate varies across three microclimate gradients: accumulated degree-days, growing-season soil moisture, and days of snow cover.Range-wide variation in demographic rates was best explained by the combined influence of multiple microclimatic variables. Different pairs of demographic rates exhibited both similar and inverse responses to the same microclimatic gradient, and the microclimatic effects often varied with plant size. These responses resulted in range-wide projected population persistence, with no declining populations at either elevational range edge or at the extremes of the microclimate gradients.The complex relationships between topography, microclimate and demography suggest that populations across a species' range may have unique demographic pathways to stable population dynamics.

Plant distributions are strongly influenced by both climate and topography. In an analysis of geographic and topographic distributions for selected tree species in California, we found that tree populations are increasingly restricted to extreme topographic positions as they approach the edge of their geographic ranges, occupying cooler, pole-facing slopes (at the warm and dry edge) and warmer, equator-facing slopes (at the cool and moist edge). At a local scale, species distributions across topographic gradients also correlate with species geographic ranges (species that occupy cooler locations within the landscape have cooler, moister geographic distributions, and vice versa). Model outputs indicated that species found on pole-facing slopes and equator-facing slopes will experience population declines and population increases, respectively, in response to a warmer and drier future. As such, tree species occupying cooler landscape locations, which are viewed as refugia in some contexts, may be most threatened by anthropogenic climate change.