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.     

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.     

Post-fire resprouting of shrubby species in Mediterranean maquis

The vegetative resprouting of mediterranean maquis shrubby species was examined eight years after fire. Post-fire regeneration occurs through the resprouting of stumps. All species (Quercus ilex L., Phillyrea latifolia L., Arbutus unedo L., Erica arborea L., Pistacia lentiscus L.) show a rapid growth in the first years after fire, and a decrease already from 4^th–5^th year.All the species survived the fire and reconstituted a community similar to that unburned in a relatively short time span.     

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.     

Shoot regeneration after fire or freezing temperatures and its relation to plant life-form for some heathland species

Shoot regeneration after prescribed burning or following the freezing temperatures of winter was monitored for nineteen heathland species present in an Arctostaphyleto-Callunetum community in northeast Scotland. Species whose renewal buds were near the surface of the ground started to grow earlier in the spring than species with renewal buds above the surface, but grouping species according to the position of their renewal bud (i.e. their life-form) did not account for all of the interspecific variation apparent. In the case of shoot regeneration after fire, species whose renewal buds were destroyed by fire because they were above-ground started to regenerate about the same time as species with belowground buds, protected from fire, but reached their maximum frequency of occurrence later. Grouping species by life-form was of limited value as a means of interpreting this interspecific variation in the timing of shoot regeneration after fire. It would be unwise to use plant life-form as the sole basis for interpreting or predicting a species' response to temperature stress when extreme temperatures occur regularly, as they do in heathland. The possible use of other plant traits to interpret and predict interspecific variation in the regeneration rate of heathland plants is discussed.Nomenclature follows Tutin et al. (1964–1980) for vascular plants. Acknowledgements. The Nature Conservancy Council and Mr J. J. Humphries kindly allowed Dinnet Moor to be used for the work presented here. One of us (RJR) received financial support for field work from the Natural Sciences and Engineering Research Council of Canada.     

Biogeographical network analysis of plant species distribution in the Mediterranean region

The delimitation of bioregions helps to understand historical and ecological drivers of species distribution. In this work, we performed a network analysis of the spatial distribution patterns of plants in south of France (Languedoc‐Roussillon and Provence‐Alpes‐Côte d'Azur) to analyze the biogeographical structure of the French Mediterranean flora at different scales. We used a network approach to identify and characterize biogeographical regions, based on a large database containing 2.5 million of geolocalized plant records corresponding to more than 3,500 plant species. This methodology is performed following five steps, from the biogeographical bipartite network construction to the identification of biogeographical regions under the form of spatial network communities, the analysis of their interactions, and the identification of clusters of plant species based on the species contribution to the biogeographical regions. First, we identified two sub‐networks that distinguish Mediterranean and temperate biota. Then, we separated eight statistically significant bioregions that present a complex spatial structure. Some of them are spatially well delimited and match with particular geological entities. On the other hand, fuzzy transitions arise between adjacent bioregions that share a common geological setting, but are spread along a climatic gradient. The proposed network approach illustrates the biogeographical structure of the flora in southern France and provides precise insights into the relationships between bioregions. This approach sheds light on ecological drivers shaping the distribution of Mediterranean biota: The interplay between a climatic gradient and geological substrate shapes biodiversity patterns. Finally, this work exemplifies why fragmented distributions are common in the Mediterranean region, isolating groups of species that share a similar eco‐evolutionary history.     

Germination activity of smoke residues in soils following a fire

A simple and rapid bioassay was implemented to detect the germination activity of extracts from soils in pre/post-burn conditions. Soil samples taken from burnt, unburnt and adjacent plots at depths of 0–2, 2–4, 4–6 and 6–8 cm before and after burning mesic grassland in South Africa were analysed for germination activity over an eight-week period. Soil samples were extracted using dichloromethane and bioassayed using Grand Rapids lettuce (Lactuca sativa L.) achenes (seeds). The Grand Rapids lettuce seeds exhibited greater germination percentages when treated with extracts from burnt soil compared to the other plots. The magnitude of the germination activity declined with time since the burn. The Grand Rapids lettuce seeds also exhibited significantly higher germination when treated with unburnt soil extracts compared to the control (distilled water) which indicates the existence of other factors controlling germination in unburnt soil. Germination activity in the adjacent plots decreased with time. These findings indicate that the germination activity of the smoke derived from burning plant-material diffuses into the soil and its persistence declines with time. Considering that the soil seed bank contains viable seeds, at a moderate depth, and that they are initially unaffected by the heat of the fire, then smoke residues following a fire can influence the germination and recruitment of plant species that are responsive to smoke-derived compounds and are represented in the germinable soil seed bank.     

Disentangling the role of heat and smoke as germination cues in Mediterranean Basin flora

Background and Aims

The role of fire as a germination cue for Mediterranean Basin (MB) plants is still unclear. The current idea is that heat stimulates germination mainly in Cistaceae and Fabaceae and that smoke has a limited role as a post-fire germination cue, in comparison with other Mediterranean-type ecosystems (MTEs), suggesting that fire-stimulated germination is less relevant in the MB than in other MTEs. However, recent studies showed that the assembly of Mediterranean plant communities is strongly driven by post-fire germination, suggesting an important role for fire as a germination cue. We hypothesize that both heat and smoke have important effects on the different post-fire recruitment processes of MB species (e.g. level and rate of germination and initial seedling growth).

Methods

To ascertain the role of heat and smoke in the post-fire germination response of MB woody plants, a germination experiment was performed with seven heat and two smoke treatments on 30 MB woody species from seven different families, including species with water-permeable seeds and species with water-impermeable seeds.

Key Results

Heat stimulated the germination (probability and rate) of 21 species and smoke in eight species, out of the 30 species studied. In addition, six species showed enhanced initial seedling growth after the smoke treatments.

Conclusions

The results suggest that both heat and smoke are important germination cues in a wide range of MB woody species and that fire-cued germination in woody plants of the MB may be as important as in other MTEs.     

Effect of fire-derived chemicals on germination and seedling growth in Mediterranean plant species

The promoting effect of smoke-derived chemicals (e.g. karrikinolide and cyanohydrin) on germination in many plants from Mediterranean-type ecosystems such as South Africa and south-western Australia is well documented. However, very little is known about (1) the relative importance of different compounds and their possible interactive effects, (2) their role in enhancing seedling growth in wild plants, and (3) their effect on the germination of plants in the Mediterranean Basin. To fill these gaps in knowledge, we performed experiments to evaluate the effect of smoke water, karrikinolide, mandelonitrile (a cyanohydrin analogue), potassium nitrate and gibberellic acid on the germination and seedling growth of 37 species from the Mediterranean Basin. The results suggest that germination and/or seedling growth of 21 species are enhanced by at least one of the fire-derived chemicals. There were positive correlations between most of the compounds tested in terms of germination response, but synergetic and inhibitory effects were also detected. Stimulation of germination was most prominent in species with annual life cycles. Fire-derived chemicals were more effective in stimulating root growth than shoot growth. In conclusion, we provide novel evidence that the recruitment of different Mediterranean species may be enhanced by different smoke compounds, and that synergetic and inhibitory effects of chemical compounds are important in the germination ecology of plants.     

Diversity patterns of plant functional types in relation to fire regime and previous land use in Mediterranean woodlands

Abstract. We explore patterns of diversity of plant functional types (PFTs) in Mediterranean communities subjected to landscape‐scale fire disturbances in a mosaic of uncultivated and old fields stands. We use regenerative and growth form attributes to establish two sets of PFTs of perennial species living in shrublands and pine forests of NE Spain. We test the following hypotheses: 1. Fire frequency decreases regenerative PFTs diversity by negatively selecting attributes with low regenerative efficiency. 2. Fire history has more influence on regenerative than on growth form PFTs. 3. The lowest diversity of growth form PFTs will be in old fields without recent fires. We surveyed stands of different combinations of fire and land use histories. Fire history included areas without fires in the last twenty years (unburned), sites burned in 1982 (1‐burned), and sites burned in 1982 and 1994 (2‐burned). Land use histories considered terraced old fields, and uncultivated stands on stony soils. We analysed patterns of PFT abundance and diversity at the stand level, and across the landscape (among stands absolute deviations from sample medians of the relative cover of PFTs). At the stand level, fire had more influence on the diversity patterns of regenerative PFTs than on growth form PFTs. Fire decreased the diversity of regenerative PFTs, due to the elimination of the species without effective mechanisms to post‐fire regeneration. This effect was not observed across the landscape, but seeders showed more variation in stands with longer history without fire. Land use contributed to explain the diversity patterns of growth form PFT (i.e. the number of growth form PFTs was lower in uncultivated, unburned sites), but it did not influence regenerative PFTs diversity. Patterns of PFTs diversity reflect the response to ecological processes operating at the landscape level. Overall, regenerative and growth form PFTs appear to be more sensitive to the fire history than to the past land use.     

Smoke interacts with fire history to stimulate soil seed bank germination in Mediterranean woodlands

Aims Fire has important consequences on vegetation dynamics. In fire-prone areas, natural selection favors plant species, characterized by a large soil seed bank, and that their germination is stimulated by fire. Although seed germination stimulated by fire heat is common in the eastern Mediterranean Basin, only little is known about germination stimulation by smoke. We examined the interactive effect of aerosol smoke and fire history on the germinable soil seed bank (GSSB) community in eastern Mediterranean woodlands.     

Effect of high temperatures and ash on germination of ten species from gorse shrubland

Seeds of ten species of Cistaceae, Ericaceae and Poaceae were examinated to compare their responses to high temperatures and ash, simulating the direct effect of fire on germination. A variable response to these factors between families and within species from the same family was detected.In Cistaceae, heat treatment pre-sowing stimulated germination. In Ericaceae, germination was stimulated by thermic shock, but a wide range of response was detected (between all four species), from Erica ciliari, very sensitive, to Daboecia cantabrica and Calluna vulgaris which showed no significant response. The range of reaction shown by the species of this family suggest that the size of the seeds is directly related to the difference in response, at least in species with a similar structure. In Poaceae, germination was not found to be dependent upon thermic shock.The effect of ash varies in magnitude. Although it decreased the germination percentage in all species, in Avenula marginata (Poaceae) the effect was not statistically significant, whilst in Ericaceae, particularly in those showing the highest levels of germination in the control (C. vulgaris and E. umbellata), the effect is to inhibit germination completely. In respect to the effect of ash the families could be arranged in the order Ericaceae>Cistaceae>Poaceae.Complex interactions between various dormancy types and seed size could explain these results. In species which depend on physical dormancy temperature has an important role. When other dormancy type is dominant in the control of germination, the medium characteristics could have a relevant role on seed response. In addition, the seed-size could determine the threshold of tolerance to temperature.     

Rareness of certain Mediterranean ant species: fact or artifact?

Summary A checklist of the ants recorded during five years in a small urban area (400 m²) in Sant Cugat (Barcelona, Spain) is presented. 23 species nest in the area, 7 species (hypogaeic or parasitic) probably nest and 5 species nest in the vicinity; five additional species have also been recovered from the area. The rarity of some of these (Leptanilla sp.,Hypoponera abeillei (Emery),Cryptopone ochraceum (Mayr),Smithistruma tenuipilis (Emery),Epitritus argiolus Emery,Trichoscapa membranifera Emery,Chalepoxenus muellerianus (Finzi),Anergates atratulus (Schenck),Plagiolepis xene (Schenck),Lasius carniolicus Mayr) and the high number of species suggest that a) rarity in Mediterranean ant species may be explained by the insufficient sampling in time of adequate microhabitats (hypogaeic species) or by the inconspicuousness of social parasitic species, and b) that previously published species lists of ants from Mediterranean habitats might be rather incomplete.     

Germination response to fire-related factors of seeds from non-serotinous and serotinous cones

Pinus halepensis, a Mediterranean pine tree, is a partially serotinous species: individual trees of this species carry both non-serotinous and serotinous cones. Serotinous cones open mainly after fire, whereas non-serotinous cones open in absence of fire. In this study we addressed the question, whether or not this cone response is linked with the germination response of seeds to fires. Two main factors associated with fire are heating of seeds and soil pH. A combination of high heat and high pH simulates a scenario with fire, whereas low heat and low pH simulates a scenario without fire. We assessed the separate and combined effects of heat and pH on the germination rate and the percentage of germination of seeds from non-serotinous cones and two age classes of serotinous cones of P. halepensis. Heat had no effect on the percentage of germination of seeds from any of the cone types, but did positively affect the germination rates of seeds from both age-classes of serotinous cones. High pH negatively affected the germination rate of seeds from all cone types as well as the percentage of germination of seeds from non-serotinous cones. The combinations of heat and pH had different effects on the three cone types: percentage of germination and rate of germination of seeds from non-serotinous cones was higher in the combination high heat-high pH than in the combination low heat-low pH. In the combination high heat-high pH, seeds from serotinous cones germinated better than seeds from non-serotinous cones. The different germination responses of seeds from non-serotinous and serotinous cones could not be attributed to differences in cone age. Our results indicate that the cone response is linked to the germination response of the seeds in P. halepensis, with seeds from serotinous cones being more tolerant to fire related factors. This revised version was published online in June 2006 with corrections to the Cover Date.     

Taking stock: inventory of alien species in the Mediterranean sea

573 alien marine metazoan species have been recorded in the Mediterranean Sea. The present checklist is the first to present the species’ native range, presumed mode of introduction, spatial extent, and the date of the first record in each country. The majority of aliens are thermophilic species originating from the Indo-Pacific or Indian Oceans, which have entered the Mediterranean through the Suez Canal. However, the means of introduction differ greatly among the phyla, and the basins of the Mediterranean. The temporal records of the alien species reflect political crises, economic development and scientific interest in studying the phenomenon—in the past two decades on average about 10 alien species new to the Mediterranean are recorded annually. Many have established durable populations and extended their range: 125 alien species have been recorded from four or more countries. The possible impacts of regulatory instruments and environmental management options are examined. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Biogeographic patterns in Mediterranean and Macaronesian species of Saxifraga (Saxifragaceae) inferred from phylogenetic analyses of ITS sequences

A biogeographic study of Saxifraga section Saxifraga was performed based on phylogenetic analyses of ITS (internal transcribed spacer) sequences of nuclear ribosomal DNA. ITS sequences from 21 species and 31 populations were examined to identify colonization patterns for the two species of Saxifraga occurring in Macaronesia and for S. globulifera in the west Mediterranean basin. Phylogenetic analysis of the sequence data yield a single most parsimonious tree with many of the major clades well supported by bootstrap and decay values. The ITS tree provided resolution at specific and populational levels that points to two biogeographic patterns within the genus. In contrast to the molecular evidence provided by other authors for a Mediterranean origin of several Macaronesian genera of angiosperms, our results indicate that the Madeiran archipelago was colonized a single time by a species of Saxifraga originating from the Eurosiberian region. On the other hand, the molecular evidence also suggests that populations of S. globulifera from North Africa have been isolated for a long time from populations occurring in the Iberian Peninsula, and that the endemic S. reuteriana has evolved from the Iberian populations of S. globulifera. The Mediterranean Sea has probably been an effective isolating barrier for some plant groups that occur in Europe and North Africa.     

The effects of fire on communities,guilds and species of breeding birds in burnt and control pinewoods in central Italy

Breeding bird communities in burnt and unburnt residual pinewoods were studied over 3 years by line-transect method, following a catastrophic fire event in Castelfusano (Rome, Central Italy; July 2000). We applied bootstrap procedures to evaluate whether the observed data were true or just produced by chance, and then examined the emerging patterns at three levels: community, guild and species levels. At the community level, fire acted on breeding bird communities by altering especially the total abundance patterns: the species abundance decreased in the burnt pinewood compared to the residual one, but other parameters were not significantly affected by fire. As a consequence of fire, the destruction and structural simplification of the canopy and shrubby component, as well as the increase of edge habitat and patchiness at landscape scale, induced a turnover in species between pinewoods. Species turnover was higher at the burnt than at the residual pinewoods, during all the 3 years of study. At the guild level, the forest species decreased strongly in terms of richness and abundance in the burnt pinewoods, contrary to the edge and open habitat species which increased in terms of richness, abundance and evenness. Edge species showed the highest turnover in burnt pinewood during the whole period of study. At species level, after an a priori subdivision (based on bibliographic search) of the various species in two ecological guilds (forest versus edge species), it was found that an a posteriori statistical analysis confirmed the expected trend, i.e. that the species which decreased significantly in burnt pinewood were essentially the forest species, whereas the species which increased were essentially the edge/open habitat ones. Overall, in order to investigate the effects of fire catastrophes on birds, the guild approach seems more exhaustive than the taxonomic community approach, where intrinsic confounding trends are present.     

Structure of ground-foraging ant assemblages in relation to land-use change in the northwestern Mediterranean region

The abandonment by humans of marginal and less productive zones signifies an important change in land use in North Mediterranean agroecosystems. Human perturbations have led to a highly diversified landscape, with a mosaic made up of patches of land at different stages of succession, from cultivated fields to closed forest. Our aim here is to characterize ant assemblages and their functional groups in response to these land-use changes. This progressive abandonment results in an initial increase in ant richness and abundance, which can reach high levels if the succession proceeds as far as woodland. In terms of the ant functional groups, this land-use change implies: (1) the appearance of Subordinate Camponotini; (2) an increase in Generalized Myrmicinae, Cryptics and Cold-climate specialists in terms of ant species richness, overall abundance and, for Generalized Myrmicinae and Cryptics, an increase in abundance percentage; (3) a decrease in percentage abundance of Opportunists; (4) a progressive decrease in species richness as well as overall and percentage abundance of Hot-climate Specialists throughout the transformation from crops to woodlands; and (5) an initial increase of Dominant Dolichoderinae followed by a decrease in ant species richness, overall abundance and percentage abundance. Using the ant functional group approach for the clearly separate stages of the regeneration process is a promising method for comparing responses of ant communities to human land use.     

The effect of stand age and microhabitat on soil seed banks in Mediterranean Aleppo pine forests after fire

Soil samples from three microhabitats (gaps, beneath shrubs and beneath trees) in five stands of various post-fire ages (6–55 years) were collected in an east Mediterranean Aleppo pine Pinus halepensis forest. Total germinable seed bank densities varied between 300 and 1300 seeds per m². Herbaceous taxa were the major constituents of the germinable seed bank in gaps, regardless of stand age. Perennials were the major components beneath shrubs in all stands except the youngest stand where herbaceous species were the major components in all microhabitats. Important tree and shrub species (e.g., Pinus halepensis, Quercus calliprinos, Pistacia lentiscus, Phillyrea latifolia) of the mature pine forest were not an important component of the soil seed bank and therefore, little resemblance was observed between the above-ground plant species composition and soil seed bank composition. This is consistent with the fact that these species regenerate by resprouting rather than by germination from the seed bank. Both microhabitats and forest-stands, which were of different ages, contributed to the variation in taxa richness, germinable seed density and diversity among samples. The effect of small-scale spatial heterogeneity (among microhabitats) was much more pronounced. In contrast to other studies, species richness, species diversity, and density of seed banks did not decrease with post-fire age. Moreover, stand age was a poor predictor for these attributes of the soil seed bank in an Aleppo pine forest. The heterogeneity plays an important role in conservation and management of this ecosystem.