Habitat specialization through germination cueing: a comparative study of herbs from forests and open habitats.

BACKGROUND AND AIMS This study examined the adaptive association between seed germination ecology and specialization to either forest or open habitats across a range of evolutionary lineages of seed plants, in order to test the hypotheses that (1) species' specialization to open vs. shaded habitats is consistently accompanied by specialization in their regeneration niche; and (2) species are thereby adapted to utilize different windows of opportunity in time (season) and space (habitat). METHODS Seed germination response to temperature, light and stratification was tested for 17 congeneric pairs, each consisting of one forest species and one open-habitat species. A factorial design was used with temperature levels and diurnal temperature variation (10 °C constant, 15-5 °C fluctuating, 20 °C constant, 25-15 °C fluctuating), and two light levels (light and darkness) and a cold stratification treatment. The congeneric species pair design took phylogenetic dependence into account. KEY RESULTS Species from open habitats germinated better at high temperatures, whereas forest species performed equally well at low and high temperatures. Forest species tended to germinate only after a period of cold stratification that could break dormancy, while species from open habitats generally germinated without cold stratification. The empirically derived germination strategies correspond quite well with establishment opportunities for forest and open-habitat plant species in nature. CONCLUSIONS Annual changes in temperature and light regime in temperate forest delimit windows of opportunity for germination and establishment. Germination strategies of forest plants are adaptations to utilize such narrow windows in time. Conversely, lack of fit between germination ecology and environment may explain why species of open habitats generally fail to establish in forests. Germination strategy should be considered an important mechanism for habitat specialization in temperate herbs to forest habitats. The findings strongly suggest that phases in the plant life cycle other than the established phase should be considered important in adaptive specialization.