Phenological Shifts Since 1830 in 29 Native Plant Species of California and Their Responses to Historical Climate Change

Climate change is affecting Mediterranean climate regions, such as California. Retrospective phenological studies are a useful tool to track biological response to these impacts through the use of herbarium-preserved specimens. We used data from more than 12,000 herbarium specimens of 29 dominant native plant species that are characteristic of 12 broadly distributed vegetation types to investigate phenological patterns in response to climate change. We analyzed the trends of four phenophases: preflowering (FBF), flowering (F), fruiting (FS) and growth (DVG), over time (from 1830 to 2023) and through changes in climate variables (from 1896 to 2023). We also examined these trends within California's 10 ecoregions. Among the four phenophases, the strongest response was found in the timing of flowering, which showed an advance in 28 species. Furthermore, 21 species showed sequencing in the advance of two or more phenophases. We highlight the advances found over temperature variables: 10 in FBF, 28 in F, 17 in FS and 18 in DVG. Diverse and less-consistent results were found for water-related variables with 15 species advancing and 11 delaying various phenophases in response to decreasing precipitation and increasing evapotranspiration. Jepson ecoregions displayed a more pronounced advance in F related to time and mean annual temperature in the three of the southern regions compared to the northern ones. This study underscores the role of temperature in driving phenological change, demonstrating how rising temperatures have predominantly advanced phenophase timing. These findings highlight potential threats, including risks of climatic, ecological, and biological imbalances.