Survival of a polymorphic species in seasonally snow-covered forests

Color polymorphism is an adaptive strategy in which a species exhibits multiple color phenotypes in a population. Often, phenotypes are variably suited to different environmental conditions which may buffer the population against variable conditions. Modern climate change is creating novel selective pressures for many species, especially in winter habitats. Few studies have quantified the benefits of polymorphism for allowing species to cope with climate-induced environmental change, particularly for species with more cryptic differences between morphs. We investigated how color polymorphism mediates selective pressures in ruffed grouse Bonasa umbellus, a winter-adapted bird species of North American forests. Ruffed grouse display phenotypic variation in plumage color, ranging from red to gray. Over five winters (2015-2022), we monitored weather conditions, habitat use, and weekly survival for 94 grouse to test whether individuals had lower survival when phenotypically mismatched with snow cover (e.g. a gray bird on a snowless landscape or a red bird in snow). Grouse phenotypically mismatched with snow cover had lower survival, but only when winter survival rates were lowest. During winters of lower overall survival, red grouse exhibited higher survival during snow-free periods, whereas gray grouse had higher survival when snow was present. We also found that open habitat negatively impacted survival, regardless of color. While the effect of phenotypic mismatch was variable among years, it was a stronger predictor of survival than land cover, suggesting that snow is an important habitat feature mediating overwinter survival. Our work offers an advancement in understanding how environmental variability affects geographic variation in and maintenance of multiple color phenotypes in seasonally-snow covered environments. Our finding that interactions between color morph and snow cover are important for conferring winter survival provides further evidence that color polymorphism may serve as a buffer against rapidly changing conditions and a pathway for persistence of polymorphic species.Keywords: Bonasa umbellus, color polymorphism, phenotypic mismatch, ruffed grouse, winter ecology, Wisconsin