Spatial variation of seedling distribution in an east Mediterranean pine woodland at the beginning of post-fire succession

Most of the area in pine woodlands is occupied by perennial seeders that regenerate from seeds in the first winter after the fire and by annuals. Control of the germination in the regenerating vegetation after wildfire is therefore a primary ecological component of the post-fire succession in this ecosystem. The aim of the study presented here was to determine the distribution of Pinus, Cistus and other plants seeds around burned Pinus halepensis trees, and to measure the conditions related to seed germination in the upper soil layers in the same locations. The study was carried out in a 50-year old planted Pinus halepensis woodland that was burned down by a wildfire in July 1995. The variation of seedbank density was determined by collecting samples under the canopies of burned trees and in a nearby open area. Pine seedbank density decreased and that of Cistus and annuals increased with increasing distance from the burned trunks. Most pine seeds were present in the ash layer while those of the other plants were in the soil. In situ germination experiments showed that seedling density decreased with distance from the burned trunks while the proportion of pines in the seedling population increased. This was a result of seedbank variation and germination inhibition by the high pH conditions caused by the ash. The establishment of sparse pine seedling under the dead tree canopies insured their rapid development without interference by other plants and played a key role in the regeneration and stability of the pine woodland community. The concomitant mass germination of the perennial seeders in the rest of the area prevented invasion by annuals.     

Regulation of the germination of Rhus coriaria, a post-fire pioneer, by heat, ash, pH, water potential and ethylene

Germination of the post-fire pioneer species Rhus coriaria , in Pinus halepensis forests on Mount Carmel, Israel, is restricted to the ash covered microsites under large burned pine trees, where the germination of other species is strongly inhibited. The aim of this study was to examine the effect of heat, ash cover, pH, water potential (Ψ _Π ) and ethylene on germination of R. coriaria seeds, in order to identify the causes of this unique germination pattern. Pre-heating to 120–140°C for 15 min was essential for the induction of seed germination. Germination percentage was increased by ash cover of 1.2 and 2.4 kg m⁻² (1 and 2 cm, respectively) but inhibited by ash cover of 6.0 kg m⁻² (5 cm). Wet pine ash from a recently burned forest had pH of 10 and Ψ _Π of −0.26 MPa. Under such conditions the germination of R. coriaria was reduced by ca 80%. On the other hand, germination was stimulated by 0.03–0.10 p.p.m. ethylene which was released by wet ash. The post-fire germination of R. coriaria is regulated by the balance between the stimulating effects of fire heat and the ethylene released by the ash, and the inhibition caused by the high pH and the low Ψ _Π caused by the ash. Its mode of dispersal by birds and these ecophysiological attributes direct germination of R. coriaria to preferred microsites under the burned canopies of large pine trees. These microsites are characterized by improved nutrition and low competition.     

pH and osmotic potential of pine ash as post-fire germination inhibitors

The dominant plant species in native pine forests in Israel ( Pinus halepensis. Cistus salviifolius and C. creticus ) regenerate from seeds after wildfires. The future structure of the regenerating forest is determined largely by the spatial distribution of the seedlings, which depends on the response of the seeds to conditions in the upper layer of the soil. The pH and osmotic potential (π) of the soil water in this layer is strongly affected by the ash which covers the burned forest floor. The effects of pH and π on the germination of the wild species noted above and the effect of pH on the germination of two crops, radish (Kaphanus sativus radicula) and oat (Avena sativa) were studied in a growth chamber. Bis-Tris propane and CAPS [3-(cyclohexylamino)-l-propanesulfonic acid] buffer solutions (pH 6–11), as well as mannitol solutions (down to –1.5 MPa) were used. The upper soil layer from a recently burned forest had pH 9 and π–0.08 MPa. Under this pH the germination of P. halepensis and C. creticus was reduced by ca half, and of C. salviifolius by 40%. Germination of radish and oats was reduced by ca 80%. Osmotic potential of –0.1 MPa did not have a significant effect on the germination of any of the species studied. We conclude that the high pH of the soil, caused by ash, is an important environmental factor that controls the regeneration of the forest plant community. Seed adaptation to the high pH may be decisive in determinig the plants' fate the fire-prone ecosystems.     

Regulation of the germination of Aleppo pine (Pinus halepensis) by nitrate, ammonium, and gibberellin, and its role in post-fire forest regeneration

Aleppo pine ( Pinus halepensis Mill.) regenerates after wildfires only by germination. This post-fire germination is controlled by temperature, light and the chemical characteristics of the ash-bed. The extremely high pH caused by the ash inhibits germination of Aleppo pine as well as of many other species. Here we report that ash extracts at neutral pH stimulate P. halepensis germination. The effects of ash extracts, nitrate, ammonium (in concentrations present in the ash) and GA₃ on the germination of P. halepensis seeds and on lipase activity at early germination were studied under controlled conditions. Sub-optimal concentrations of nitrate- and ammonium-stimulated germination and the activity of lipase in the seeds. It is proposed that nitrate increases seed sensitivity to GA₃ while ammonium directly affects lipase activity. The balance between the inhibition of germination by the high pH of ash and its stimulation by the nitrate and ammonium present in the ash determines the overall effect of the ash on post-fire germination of Aleppo pine. The pine seedlings appear in high density among the trees but because of the effects of the ash they appear in sparse stands under the projection of the burned pine canopies. These seedlings benefit from weak inter- and intra-specific competition as well as improved mineral nutrition and have good chances to grow into large trees that will form the regenerating forest; therefore, the enhancement of P. halepensis germination by nitrate and ammonium and its inhibition by high pH can be considered as ecophysiological adaptations to post-fire regeneration.