No-analog climates and shifting realized niches during the late quaternary: implications for 21st-century predictions by species distribution models

Empirically derived species distributions models (SDMs) are increasingly relied upon to forecast species vulnerabilities to future climate change. However, many of the assumptions of SDMs may be violated when they are used to project species distributions across significant climate change events. In particular, SDM's in theory assume stable fundamental niches, but in practice, they assume stable realized niches. The assumption of a fixed realized niche relative to climate variables remains unlikely for various reasons, particularly if novel future climates open up currently unavailable portions of species fundamental niches. To demonstrate this effect, we compare the climate distributions for fossil-pollen data from 21 to 15 ka bp (relying on paleoclimate simulations) when communities and climates with no modern analog were common across North America to observed modern pollen assemblages. We test how well SDMs are able to project 20th century pollen-based taxon distributions with models calibrated using data from 21 to 15 ka. We find that taxa which were abundant in areas with no-analog late glacial climates, such as Fraxinus, Ostrya/Carpinus and Ulmus, substantially shifted their realized niches from the late glacial period to present. SDMs for these taxa had low predictive accuracy when projected to modern climates despite demonstrating high predictive accuracy for late glacial pollen distributions. For other taxa, e.g. Quercus, Picea, Pinus strobus, had relatively stable realized niches and models for these taxa tended to have higher predictive accuracy when projected to present. Our findings reinforce the point that a realized niche at any one time often represents only a subset of the climate conditions in which a taxon can persist. Projections from SDMs into future climate conditions that are based solely on contemporary realized distributions are potentially misleading for assessing the vulnerability of species to future climate change.