Germination and survival of water-impermeable seeds of 11 species of Geraniaceae and Malvaceae were monitored during dry storage and during burial in soil for up to 2.5 years. During dry storage, seeds of annual Geraniaceae became permeable and also lost their physiological dormancy. However, during burial in natural conditions, most seeds remained impermeable and viable, with no seasonal change in germination capacity. Germination in only one species (Geranium robertianum) was enhanced by daily alternating temperatures when seeds were exhumed in spring. Drying of exhumed seeds broke physical dormancy. Seeds of the perennial Geranium pratense gradually became permeable in a prolonged germination test of 31 weeks. Most seeds of Malva remained impermeable during dry storage. Buried seeds gradually germinated in situ, and exhumed seeds had a low germination capacity in all seasons. We concluded that dormancy of hard seeds in natural conditions may be broken by drying during summer, by specific temperature regimes or by gradual softening of the seed coat, ensuring the spread of germination over many seasons.
