Resprouting vigour of two mediterranean shrub species after experimental fire treatments

Experimental fire treatments were carried out by applying a propane torch flame to individual stumps ofArbutus unedo andErica arborea, two dominant ericaceous shrubs living in relatively moist maquis of the western Mediterranean Basin. No mortality was observed in either species. Individual plant size, measured as the individual stump area, was the most important factor determining both number and biomass of resprouts for all fire treatments. The number of resprouts at 3 months was less at the higher temperature, but no difference could not be detected at 18 months. Duration of flame application had no effect on resprouting success. This effect was not statistically significant 18 months after treatments were applied. Biomass of resprouts was not clearly affected by different fire treatments either 3 months or 18 months after the start of the resprouting process.     

Effects of cutting and rainfall pattern on resprouting vigour and growth of Erica arborea L.

Abstract. This study analyses the effects of recurrent cutting and rainfall pattern on sprout and genet growth in the resprouting shrub Erica arborea. Three different intervals between consecutive clear-cutting events were applied: 30, 5 and 2 yr. Above-ground genet biomass after 2 yr of treatment application was correlated with burl size and decreased with increasing recurrence of cutting, indicating that more than 5 yr were needed to re-establish the resprouting potential of genets. The effect of rainfall on growth was evaluated through the correlations between sprout RGR and several variables related to the rainfall pattern at different regeneration stages. The effect of the rainfall pattern on sprout RGR was dependent on the regeneration age, suggesting that water availability becomes more limiting after two years of regrowth.     

Resprouting of Quercus suber in NE Spain after fire

Abstract. Many Mediterranean species have evolved strategies that allow them to survive periodic wildfires. Quercus suber trees resprout after fire, some from stem buds and others from basal buds only. In the former case the canopy recovers quickly. In the latter case the stem dies but the tree survives and regrows from basal sprouts. The probability of stem death and the degree of height recovery were studied after a fire in a Q. suber forest in NE Spain using logistic regression analysis. The results suggest that most trees survive after fire; the probability of stem death is negatively related to tree diameter; and recovery is positively related to tree diameter and to bark thickness. Implication for management and conservation of cork-oak forests are discussed.     

Effects of Rosmarinus officinalis neighbors on resprouting of Erica multiflora individuals

To study the effects of competition in Mediterranean shrubland regeneration following disturbance, we used a neighborhood approach to assess the influence of mature Rosmarinus officinalis neighbors on the resprouting of Erica multiflora individuals after clipping. Sprout biomass of target plants 2 years after clipping was regressed against various measures of neighbor abundance within a 2 m radius around target E. multiflora individuals in which all vegetation except R. officinalis had been removed. The largest single influence on the biomass of sprouts produced was the previous biomass of the resprouting plant. The abundance of R. officinalis neighbors had a weak but detectable effect on resprouting of E. multiflora. Abundance of neighbors within 60 cm from target plants was the best predictor of regrowth. At this distance, two simple measures of neighbor abundance within the neighborhood, the number of neighbors and the sum of their heights, were significant in accounting for variation in resprouted biomass. None of the combinations of neighbor variables performed significantly better than single variables. The best models accounted for around 24 percent of the variation in resprout biomass. As in other studies, angular dispersion of neighbors never had a significant effect on performance of target plants. The weak but significant response of resprouting to variation in R. officinalis abundance suggests that the intensity of competition in the experiment was low because of the removal of other species.     

Resprouting in the Mediterranean‐type shrub Erica australis afffected by soil resource availability

Abstract. Soil resource availability may affect plant regeneration by resprouting in disturbed environments directly, by affecting plant growth rates, or indirectly by determining allocation to storage in the resprouting organs. Allocation to storage may be higher in stressful, low resource‐supply soils, but under such conditions plant growth rates may be lower. These factors could act in opposite directions leading to poorly known effects on resprouting. This paper analyses the role played by soil resources in the production and growth of resprouts after removal of above‐ground plant tissues in the Mediterranean shrub Erica australis. At 13 sites, differing in substrate, we cut the base of the stems of six plants of E. australis and allowed them to resprout and grow for two years. Soils were chemically analysed and plant water potential measured during the summer at all sites to characterize soil resource availability. We used stepwise regression analysis to determine the relationships between the resprouting response [mean site values of the number of resprouts (RN), maximum length (RML) and biomass (RB)] and soil nutrient content and plant water potential at each site. During the first two years of resprouting there were statistically significant differences among sites in the variables characterizing the resprouting response. RML was always different among sites and had little relationship with lignotuber area. RN was less different among sites and was always positively correlated with lignotuber area. RB was different among sites after the two years of growth. During the first months of resprouting, RN and RML were highly and positively related to the water status of the plant during summer. At later dates soil fertility variables came into play, explaining significant amounts of variance of the resprouting variables. Soil extractable cations content was the main variable accounting for RML and RB. Our results indicate that resprout growth of E. australis is positively affected by high water availability at the beginning of the resprouting response and negatively so by high soil extractable cation content at later periods. Some of these factors had previously shown to be related, with an opposite sign, to the development of a relatively larger lignotuber. Indeed, RML and RB measured in the second year of resprouting were significantly and negatively correlated with some indices of biomass allocation to the lignotuber at each site. This indicates that sites favouring allocation to the resprouting organ may not favour resprout growth.     

Sprout recruitment and self-thinning of Erica multiflora after clipping

Regrowth after clipping and the effect of local competition were studied in a natural population of Erica multiflora in a Mediterranean shrubland, by removing neighbours at 1 and 2 m around the target plants during four growing seasons. Removal of surrounding natural vegetation increased the number, the density (number of sprouts per stump area) and the biomass of the sprouts growing from clipped plants. Target plants ònly interacted with their near neighbours. Target plants had a negative relative increment in the number of sprouts per stump during the 18 months immediately following treatment, but a positive increment thereafter, which suggests that there was a constant or episodic recruitment of sprouts within the stump after clipping. Competition treatment had a non-significant effect on the negative increment of sprouts per stump. The self-thinning trajectory was different for the different competition treatments: there was an allometric negative relationship between density of sprouts and mean biomass of survivors during all sampling periods in genets without neighbours in a 1-m radius; the self-thinning trajectory of sprouts in genets without neighbours in a 2-m radius was short, a net increase in sprouts per stump area was accompanied by an increase in mean sprout biomass 30 months after clipping. During the same period, however, plants with neighbours showed a decline in both the sprout biomass and density.     

Comparative genet survival after fire in woody Mediterranean species

Summary Using data from three fires in northeastern Spain, we tested a condition necessary to support the idea that fire has been a factor in the evolution of the resprouting habit: populations of all resprouting species within a community should show high levels of genet survival after fires and show a low coefficient of variation. Species with high mean survival values were:Quercus ilex L.,Phillyrea latifolia L., andViburnum tinus L., with 88, 86 and 83% survival respectively; these groups had resprouts emerging from rootcrowns. Then followedArbutus unedo L. (75%),Pistacia lentiscus L. (73%),Erica arborea L. (77%),Erica multiflora L. (57%) andJuniperus oxycedrus L. (55%). This last group had resprouts from lignotubers or burls. These two groups also differed in the variability around the mean: the first showed a lower coefficient of variation, 6–12, and the second ranged from 19 to 26. Slope exposure had no significant influence on the process of resprouting, but soil depth did, with precipitation as a covariate. In the shallow soil category, the difference in genet survival between southern and northern exposures was 14% (71% vs. 57%); while the difference in the deep soil category was low, 5% (87% vs. 82%). There was no significant interaction. The component of variance for soils was larger than that for species-specific effects; substantial overlap of the within-species variance indicated that species responded as if they were a single hypothetical population, in which most of the variation in chances of survival was due to the soil conditions. The possession of the resprouting habit did not ensure a high performance. Hence, we find weak support for fire as a factor in the evolution of the resprouting habit.     

Interactions between mediterranean shrub species eight years after experimental fire

This paper is part of a wider study in which different combinations of species were assayed for the revegetation of experimentally burnt heathlands. In this particular case, we determined whether there was any interaction between the dominant species, Erica australis, which recovered by vegetative resprouting, and the only successful woody species sown, Cytisus sp., eight years after fire and seven years after sowing. A detailed study was carried out on each square metre of cover and height of both species in three 160 m² plots. Total cover exceeded 100% in all subplots sown with Cytisus, whereas it did not reach 65% in the others, when both species coexist together. There was an inverse correlation between the covers of both species and a positive correlation between the height of Erica australis and the cover and the height of Cytisus. There was no significant difference in Erica cover in plots sown with Cytisus and plots not sown. There could be some competition for light, but both species are capable of coexisting with high canopy overlap, for at least some time. Cytisus is expanding from the seeded subplots, suggesting that the dense cover of Erica australis does not impede the establishment of Cytisus. This revised version was published online in August 2006 with corrections to the Cover Date.     

Changing the fire management regime in the renosterveld and lowland fynbos of the Bontebok National Park

This paper evaluates the history of fire management in the Bontebok National Park (3435 ha) over a period of almost four decades. A GIS database was compiled of all fires between 1972 and 2009 and the fire regime was analysed in terms of the frequency, season, size and cause of fires. Since the early 1970s, short interval burning was implemented to promote grazing for bontebok, but from 2004 the fire interval was lengthened to favour plant species diversity, an increasingly urgent conservation priority for the park. In total, 43 fires were recorded, ranging in size from 9 to 1007 ha, collectively spanning 14 013 ha. The majority of fires were large (100–500 ha), with fires of >100 ha accounting for 96% of the area burnt. The overall mean fire return period (FRP) for the park was 7.2 years, which is short judged by fynbos standards. FRPs under the old and new management regimes were 6.7 and 10.9 years respectively. Under the old regime, FRPs in renosterveld and fynbos were 5.8 and 8.0 years respectively. Large parts of the park repeatedly experienced fires at immature vegetation ages resulting in the elimination of slow-maturing seed-regenerating plant species such as Protea repens. Post-fire age distribution was highly skewed towards young vegetation, with 75% of fire-prone vegetation burning at post-fire ages of ≤7 years, and <10% of fire-prone vegetation surviving beyond 10 years of age. Prescribed and accidental fires respectively accounted for 70% and 30% of the total area burnt. Prescribed burning was mostly done in March–April, and only 8% of the total area burnt, burnt outside of the ecologically acceptable fire season. This study identified areas which have been subject to ecologically appropriate and inappropriate fire return intervals, providing a basis for informed future management and research.     

Effects of nutrient availability and neighbours on shoot growth,resprouting and flowering of Erica multiflora

Abstract. To test if low soil fertility and competition limit the performance of Mediterranean shrubs, and if the effects of competition on plant performance were modified by soil fertility, we subjected shrubs of Erica multiflora to a factorial field experiment of fertilization and removal of neighbours around target plants. After 18 months of treatment, fertilization had stimulated the growth of pre-existent sprouts and biomass allocation to stems into new sprouts, but decreased the frequency of sprout flowering. Removal of neighbours increased the number and biomass of new sprouts, the probability of sprout flowering and the biomass of flowers. Fertilization slightly enhanced sprout recruitment and the probability of sprout flowering when neighbours were removed, but did not modify the other parameters of plant performance. According to our results, both low soil fertility and competition limited plant performance. Competition was slightly more intense in fertilized plants, but only in determining sprout and flowering bud stimulation.     

Drought advances spring growth phenology of the Mediterranean shrub Erica multiflora

Current climate projections predict drier and warmer conditions in the Mediterranean basin over the next century. While advanced spring growth due to warming has been described in the literature, few data are available on the effects of drought on phenology. Hence, the phenology and growth of two Mediterranean shrubs, Erica multiflora and Globularia alypum, was studied in a rainfall exclusion field experiment to simulate spring drought in a natural shrubland. We estimated the onset of growth in spring by monitoring the appearance of new stems, and the end of growth in summer by following the elongation of stems. Drought treatment caused earlier onset of the spring growing season in E. multiflora, whereas no advance was observed in G. alypum. However, growth cessation was not affected in E. multiflora. Drought reduced the growth of both shrubs, as reflected in less stem elongation. The results show that a drier climate might affect not only growth but also spring phenology of some Mediterranean species. We suggest that a reduction in the cooling effect of transpiration may have analogous effects to warming and might advance the start of growth in E. multiflora, a species whose phenology has been described as warming‐sensitive. The lengthening of the growing season resulting from advanced growth did not imply higher productivity, as growth was restricted by drought.     

Diversity and stability in garrigue ecosystems after fire

Summary Vegetation dynamics after fire was studied in six communities in Bas-Languedoc (Southern France). 47 plots were observed by means of a permanent transect for ten years.In the first part, we describe floristic richness, species fugacity and the way by which, the terminal community (as defined by the last observation) appears. The dynamics of all these communities follows a simple and general model: floristic richness reaches its maximum during the first two years after a fire, then decreases and becomes stable. Fugacity follows a similar model, whereas the mergence of the terminal community is rapid: one year after fire 70% of the plots have already acquired 75% of the species of the terminal community. There is no succession (in the general sense of the word), but a progressive reappearance of the species belonging to the original community.In the second part, we study floristic similarities between our plots and corresponding associations as described in literature. It appears that after a fire the floristic diversity of the landscape remains high; while the communities rapidly reach a relative maturity.In the study area fire seems to be a rather superficial phenomenon; it does not lead to an important modification of the community dynamics, because probably the most frequent species in Bas-Languedoc developed adaptations to withstand fire.     

Local survival after fire in Mediterranean shrublands: combining capture-recapture data over several bird species

Post-disturbance survival is a key factor in the onset of secondary successions. Here we analyse capture-recapture data from two before/after disturbance studies to estimate the effect of fire on local bird survival. Analyses of six bird species at two Mediterranean shrubland sites were combined using a meta-analysis approach. Two warblers, Sylvia undata and S. melanocephala, were studied at one site altered by prescribed burning, and five passerines (Luscinia megarhynchos, Turdus merula, Parus major, P. caeruleus and S. melanocephala) at one site disturbed by wildfire. Based on the combined analysis, annual survival probability significantly decreased from 0.49 to 0.18 (i.e. a 64% decline) after the fire. Our results further suggest a trend for a higher decrease in annual survival associated with wildfire (–72%, from 0.51 to 0.14) than with prescribed burning (–35%, from 0.41 to 0.27), although this should be properly tested with a specific experimental design. In S. undata, a decline in survival in the long-term cannot account for the drop in density observed the first spring after fire. We suggest that a decrease in recruitment rate and an increase in the proportion of non-breeders immediately after the fire may contribute more strongly to the decline in the breeding population. Our results tend to support the idea that bird populations may respond to moderate disturbances with noticeable time lags, because of individual site tenacity.     

The influence of fire on the phenological behaviour of Mediterranean plant species in Bas-Languedoc (southern France)

Fire is an ancient ecological factor influencing the Mediterranean vegetation of southern France. The study was carried out on three areas to determine the phenological behaviour of plants with regard to fire. First we studied the flowering responses of perennials in relation to the time since fire: in a Quercus coccifera garrigue most species flower during the year following burning. In comparing species by species between burned and unburned areas most species did not show major differences in the phenological stages. However, fire did increase the number of inflorescences of grasses. A phenological synthesis showed that differences at the community level existed for the flowering stages between the burned areas and the unburned control sites during the first and second years following fire. The growth of some woody species was also studied; the elongation and growth of the plants were biggest during the first or second year after fire. The lack of differences in phenological response between burned and unburned plants may be an adaptive trait to fire.     

Seed banks of Erica australis and Calluna vulgaris in a heathland subjected to experimental fire

Abstract. Soil samples were collected before and after an experimental fire on a heathland in the province of León (Spain). The seed banks were assessed by counting the numbers of seedlings appearing of Erica australis and Calluna vulgaris. A total of 2285 germinable seeds/m² before burning and 1177 germinable seeds/m² after the fire were estimated for Erica australis, and 90 and 690 germinable seeds/m², respectively, for Calluna vulgaris. After the fire the number of germinable seeds/m² of Erica australis had decreased, whilst there was a considerable increase in the number of germinable seeds/m² of Calluna vulgaris. Neither seedlings nor sprouts of Calluna vulgaris had appeared in the field plots 10 months after the fire. Erica australis did recover mainly by sprouting after fire.     

Fire regenerative syndromes of forest woody species across fire and climatic gradients

Community resilience after fire is determined by species’ ability to regenerate through two main mechanisms growth of new sprouts (resprouter species) and germination from surviving seed banks or from seeds arriving from neighbouring populations (seeder species). Both the mechanisms are present in Mediterranean communities. The occurrence of both the types in a community depends on fire history and the bio-geographical history determining the available species pool. Regenerative traits also covary with other functional attributes associated with resource acquisition and stress tolerance. As post-fire regenerative responses can be related to various ecological factors other than fire, we tested the hypothesis of a different proportional representation of post-fire regenerative syndromes in forest woody species along a climatic gradient in Catalonia (NE Spain) ranging from Mediterranean to temperate-boreal climates. Specifically, we expected seeder species to become less common with colder and moister conditions while resprouters would not be so influenced by the climatic gradient. We also tested the hypothesis of change in the relative abundance of regenerative syndromes in relation to recent fire history. We analysed a large database obtained from extensive forestry surveys and remote sensing fire records. After correction for spatial autocorrelation, we found an increase in the proportion of seeder species under more Mediterranean conditions and a decrease in fire-sensitive species (with no efficient mechanisms of post-fire recovery) in moister conditions. Resprouter species were similarly present across the whole gradient. A similar pattern was observed after excluding recently burnt plots. Therefore, post-fire regenerative syndromes segregate along the climatic gradient. Recent fires reduced the occurrence of fire-sensitive species and increased the proportion of seeder species. No significant effect was observed on resprouter species. Fire has a sorting effect, shaping the occurrence of species with different regenerative traits. Overall, fire seems to explain better the variability of the proportion of fire-sensitive species and climate the variability of seeder species. In addition, other factors (forestry practices and the covariation between regenerative and functional attributes) are likely to contribute to the regional pattern of regenerative syndromes.     

Germination behaviour of 14 Mediterranean species in relation to fire factors: smoke and heat

Fire is an ecological factor that has been present in the ecosystems of the Mediterranean region for thousands of years. Our study was undertaken to acquire knowledge of the effect of fire on the germination of Mediterranean species. We used high temperatures (up to 60°C) and smoke to determine the effect of these factors on the germination of species from the Mediterranean region. The species selected are characteristic of the central Mediterranean basin and are representative of both woody and herbaceous species: Rhamnus alaternus L., Cistus albidus L., Cistus monspeliensis L., Fumana ericoides (Cav.) Gand., Rosmarinus officinalis L., Melica ciliata L., Avena sterilis L., Bituminaria bituminosa (L.) C.H. Stirt., Anthyllis vulneraria L., Coronilla glauca L., Argyrolobium zanonii (Turra) P.W. Balland, Emerus major Mill., Genista scorpius (L.) D.C. and Spartium junceum L. The seeds were collected in Mediterranean shrubland (8) and woodland (6) ecosystems, around Montpellier, France (24°45′N and 3°50′E). Ten treatments were tested: a control, three smoke treatments and six heat treatments. The average germination level (germination percentage) and the average T₅₀ rates (time taken to reach 50% of germination) were calculated. The smoke and heat act in a different way on each of the species. The smoke enhanced the germination of two species, whilst moderate heat increased germination in all of the species excepting R. officinalis, F. ericoides, A. sterilis, A. vulneraria, and G. scorpius. Germination was fastest in M. ciliata and S. junceum and slowest in A. sterilis, E. major and C. albidus. The cues did not significantly affect the rate of germination. Fire modified the germination response of 12 of the 14 species studied.