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Fire and high summer soil temperatures can break physical seed dormancy in Mediterranean fire-prone ecosystems. Their independent effect is somewhat recognized but both factors may act together with a synergistic effect yet unknown. This study aims to determine the isolated and combined effects of fire and summer temperatures on the release of physical seed dormancy in Cistaceae species. Fire and summer temperature treatments were applied in a factorial experiment to seeds of 12 species of Cistaceae. Seeds previously exposed or not to a heat shock (fire simulation) were kept for 1 or 2 months at constant or alternating temperatures (summer temperatures simulation). Additionally, I compared the effect of exposing the seeds to a heat shock before or after they had been subjected to the summer temperatures. Heat shock increased germination of all species, but summer temperatures produced different results. When seeds were exposed to summer temperatures after heat shock, germination decreased. This negative effect disappeared when heat shock was simulated at the end of the summer temperatures. Fire and summer temperatures modulate timing of germination in Cistaceae with a joint control on post-fire regeneration. Cycling of sensitivity to physical dormancy release may be the mechanism to explain this fine-tuning, which would ensure germination when environmental conditions are suitable for growth. These results contribute to our understanding of vegetation dynamics and postfire regeneration in Mediterranean ecosystems.

Fire and high summer soil temperatures can break physical seed dormancy in Mediterranean fire-prone ecosystems. Their independent effect is somewhat recognized but both factors may act together with a synergistic effect yet unknown. This study aims to determine the isolated and combined effects of fire and summer temperatures on the release of physical seed dormancy in Cistaceae species. Fire and summer temperature treatments were applied in a factorial experiment to seeds of 12 species of Cistaceae. Seeds previously exposed or not to a heat shock (fire simulation) were kept for 1 or 2 months at constant or alternating temperatures (summer temperatures simulation). Additionally, I compared the effect of exposing the seeds to a heat shock before or after they had been subjected to the summer temperatures. Heat shock increased germination of all species, but summer temperatures produced different results. When seeds were exposed to summer temperatures after heat shock, germination decreased. This negative effect disappeared when heat shock was simulated at the end of the summer temperatures. Fire and summer temperatures modulate timing of germination in Cistaceae with a joint control on post-fire regeneration. Cycling of sensitivity to physical dormancy release may be the mechanism to explain this fine-tuning, which would ensure germination when environmental conditions are suitable for growth. These results contribute to our understanding of vegetation dynamics and postfire regeneration in Mediterranean ecosystems.

In fire-prone environments, the "event-dependent hypothesis" states that plant population changes are driven by the unique set of conditions of a fire (e.g., fire season, climate). Climate variability, in particular changes in rainfall patterns, can be most important for seeder species, since they must regenerate after fire from seeds, and for Mediterranean shrublands, given the high yearly variability of rainfall in these ecosystems. Yet, the role of rainfall variability and its interaction with fire characteristics (e.g., fire season) on plant populations has received little attention. Here we investigated the changes in seedling emergence and recruitment of three seeder species (Cistus ladanifer, Erica umbellata and Rosmarinus officinalis) after fires lit during three different years and at two times during the fire season (early and late in the fire season) to account for potential changes in the soil seed-bank during the year. Three plots were burned at each season, for a total of 18 plots burned during the three years. After fire, emerged seedlings were tallied, tagged and monitored during three years (two the last burning year). Rainfall during the study period was rather variable, and in some years was well below average. Seedling emergence after fire varied by a factor of 3 to 10, depending on the species and on the burning year. The bulk of seedling emergence occurred in the first year after fire, and seedling recruitment at the end of the study period was tightly correlated with this early emergence. Seedling emergence in E umbellata and R officinalis, but not in C ladanifer, were correlated with precipitation in the fall and winter immediately after the fire, being E umbellata most sensitive to low rainfall. Fire season was generally not an important factor in controlling emergence and recruitment. We discuss how projected changes in rainfall patterns with global warming can alter the balance of species in this shrubland, and can drive some species to near local extinction.

Soil seed banks play a major role in the post-fire regeneration of Mediterranean shrublands. They vary throughout the year in species composition, abundance, and readiness to germinate. After fire, germination occurs mainly during the following fall to spring. Time of germination can determine recruitment success. It is unclear what factors control post-fire germination and its timing. We tested the effects of season and fire on the readily germinable soil seed bank of a seeder-dominated shrubland. Plots were burned early and late in the summer season (ES, LS). Soil samples were collected before and after fire, and germinated in a chamber simulating successively autumn, winter, and spring conditions. Samples were kept moistened at all times. Fire intensity was similar between ES and LS. Several species of Cistus and herbs, mostly annuals, were dominant. Most germination occurred during the simulated-autumn period, with little subsequent germination during the following two periods. Germination speed (T 50) during simulated-autumn was similar for shrubs and herbs, and independent of season or fire. Germination was lower for two shrubs (Rosmarinus officinalis, Cistus salvifolius) and higher for herbaceous dicots in LS than in ES soils. Fire reduced monocots and enhanced Cistus. Germination period significantly interacted with fire and season in some groups or species, altering the simulated-autumn germination peak. We demonstrate that the seed bank can germinate swiftly under simulated-autumn conditions. Hence, water availability is the main controlling factor of germination. Fire season differentially affected some species or groups, and could affect the post-fire regeneration.

Little is known about consequences of the increasing global temperatures on seed germination and even less about the seed germination response of species with different geographical distribution ranges. In the present work, the effect of incubation temperature on seed germination of 31 species from the Iberian Peninsula is analyzed in relation to their geographical distribution range. We tested whether plants with different geographical distribution ranges (Iberian Peninsula endemics, Mediterranean, widely distributed species), respond similarly to incubation temperature using cross-species analysis as well as taking into account phylogenetic relationships among species. Seeds were incubated at 10°C, 15°C, 20°C and 25°C for 6 weeks in darkness. Distribution range appeared as a determinant of germination response to the temperature treatment independent of phylogeny. Widely distributed species were less sensitive than endemic or Mediterranean species to incubation temperatures, which may represent an advantage to colonize new areas or to tolerate changes in the environment. Although their ability to avoid germination during the warm and dry season might be favorable for endemic species in some situations, our results also suggest that under a future of temperature increase imposed by climate change, endemic plant species may be more vulnerable than widely distributed species.

The effects of cold stratification on seed germination of 58 species from a Mediterranean area in central–eastern Spain were studied. The area is subject to fires and has a very continental climate, with long winters (187 frost days during the year). Differences in germination among species taking into account their phylogeny were studied by dividing them according to their life form (chamaephytes, hemicryptophytes), post-fire regeneration strategy (non-sprouters, sprouters), and geographical distribution range (Iberian Peninsula endemics, Mediterranean, widely distributed species). Treated seeds were stratified at 5 °C for 4 weeks, then incubated, along with the control seeds, at 15 °C for 6 weeks. Most species were not affected by cold stratification, and in the cases where they were, germination was generally lower after stratification, not higher. In general, germination percentages were low, which shows that stratification by itself was not able to break seed dormancy. Functional groups diff...

Post-fire conditions are characterized by enhanced light and the availability of nitrogenous compounds in the soil. It is not known, however, to what extent light or nitrogenous compounds control the germination response of species growing in burned areas and, in particular, whether functional groups of plants differ in their response. The germination response to light and nitrate was tested for 53 species representative of the flora of a Mediterranean recently burned area in Central-Eastern Spain. Differences in germination among species, with and without taking into account their phylogeny, were studied by classifying them according to their life-form (chamaephytes, hemicryptophytes), regeneration strategy (non-sprouters, sprouters) and geographical distribution range (Iberian Peninsula endemics, Mediterranean, widely distributed species). The overall germination mean was not affected by any of the two treatments. There were statistically significant interactions between species and the two treatments. That is, not all species were equally affected, and about 30% of the species were significantly affected by light (half of them positively and the other half negatively) and 25% by nitrate (most of them positively). Species response was related to functional groups. Light response (stimulation vs. non-stimulation) was associated to life-form, regeneration strategy and distribution range. Hemicryptophytes, sprouters and widely distributed species were positively affected by light. No evidence of such an association for nitrate was found. No statistically significant effects of light and nitrate on the mean germination of the various groups (life-form, regeneration strategy, distribution range) were found. Moreover, significant interactions emerged between light and nitrate for all three groups. In summary, the studied set of plants appears to be non-dependent on factors that may change with fire, such as increased light and soil nitrate, for germination. Nonetheless, some species and groups will be affected by such changes. That means that fire will modify the relative balance of germination among species and functional groups. Due to the particular sensitivity of Iberian Peninsula endemics to light, a factor that significantly changes with fire, these species may be at risk under the current fire regime.

Many plants from Mediterranean-type ecosystems show enhanced germination when exposed to heat and other fire-related cues. Whether Mediterranean-type plants are fire recruiters and need fire to germinate is subject of discussion. In the present work, we tested the above hypothesis by subjecting seeds from 57 species representative of a recently burned plant community to various heat pulses. Differences in germination among species, taking into account their phylogeny, were studied by dividing them according to their life-form (chamaephytes, hemicryptophytes), regeneration strategy (non-sprouters, sprouters) and geographical distribution range (Iberian Peninsula endemics, Mediterranean, widely distributed species). Seeds were exposed to 80, 100 and 120 °C during 10 min, and then, incubated at 17.5 °C for 6 weeks in darkness. Maximum germination was reached at 80 °C and control treatments and minimum germination was at 120 °C. Most species were tolerant to heat-shocks, only a few of them were stimulated by heat, and around 20% suffered a reduction in germination with heat. The germination response to heat-shock differed according to regenerative strategy and distribution range, but not to life-form. These results were supported independently of phylogeny. As expected, sprouters and Iberian endemics were more sensitive to heat-shock than non-sprouters and widely distributed species, respectively. As a conclusion, the studied plant group appears to exhibit resistance to heat but not dependence on it for germination. Therefore, recruitment could be high after low intensity fires, but might be significantly reduced for most species after more intense fires.

This work documents the effect of a common, long-term fire retardant chemical, Fire-Trol 934, on seed viability and germination of 36 plant species growing in a burned Mediterranean area, covering different life-form types, regenerative strategies and distribution ranges. Seeds were subjected to four treatments: control, and application of Fire-Trol 934 at concentrations of 0.02, 0.2 and 2%. Fire-Trol 934 significantly decreased both seed viability and germination in the group of species studied, which suggests that Fire-Trol 934 may be toxic for seeds, at least when applied at high concentrations. Whereas seed viability generally showed a progressive decrease with increased Fire-Trol 934 concentration, germination percentages generally increased when intermediate Fire-Trol 934 concentrations were used, but tended to be drastically reduced when seeds were exposed to the highest (2%) concentration. The reduction observed in germination at the highest Fire-Trol 934 concentration was greater than that observed in viability, which suggests that the effect of Fire-Trol 934 on seeds may not be lasting. Little differences in the response to Fire-Trol 934 emerged among plant groups, all of which followed the general tendency described above.