“…Some previous studies have suggested that cone variation, such as cone size or thickness of scales in Pinus , should have adaptive significance as a means to maintain seed production and may be the responses to protect against snowfall or low temperature (Farjon & Styles, 1997; Gil et al, 2002; Linhart, 1978). As cone development carries a high cost to achieve seed production (Despland & Houle, 1997; Goubitz, Werger, Shmida, & Ne'eman, 2002; Iwaizumi & Takahashi, 2012), producing relatively highly reproductive cones is important for maximizing reproductive success in consideration of the development cost, particularly in environments with low temperature, less solar irradiance or less sunshine (Iwaizumi, Ohtani, & Takahashi, 2019). If environmental factors impact cone morphology, geographical variation linked to key influential climate differences is expected among populations or regions.…”