Variation in vegetative and flowering phenology in a forest herb caused by environmental heterogeneity

Timing of seasonal plant development can affect biotic interactions and plant fitness. Phenology is governed largely by temperature and may therefore be affected by global climate warming, making this an important area of research. Several factors in addition to temperature may cause differences in phenology. We studied the influence of local environment, plant size, and reproductive effort on shoot emergence and flowering time of 290 individuals of Actaea spicata (Ranunculaceae), distributed among 25 plots in four populations. We used multiple regression and structural equation models (SEM) to study causal relationships. Among plots, soil temperature and canopy cover explained 63% of the variation in shoot emergence. Soil temperature, slope, and canopy cover together explained 83% of the variation in flowering time. Within plots, small plants on steep south-facing slopes with high soil potassium concentrations emerged earlier in the year. Plants emerging earlier flowered earlier, but no environmental factors affected flowering time directly. We found no effects of reproductive effort. Our results support the view that flowering time of temperate forest herbs is constrained by several environmental factors acting indirectly through effects on shoot emergence time.