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.     

Plant species diversity changes in abandoned chestnut (Castanea sativa) groves in southern France

Over the last few decades, marked land use changes have taken place in many Mediterranean ecosystems. For example, many chestnut groves in France are now abandoned and have turned into 'natural' coppice stands while others are now clear-cut every 10 or 15 years for wood. Species composition and life form diversity of the plant communities have changed markedly both in the abandoned groves and in the periodically clear-cut coppice stands. What are the consequences of these changes on biodiversity at local and regional scales, and what are the implications for 'new forestry' management intended to conserve biodiversity at the same time as it optimizes productivity and profitability? To answer these questions we studied plant species diversity in the understorey strata along a successional gradient including cultivated grove; abandoned grove; young (15 years old), medium (40 years), and old (>55 years) coppice stands. The results showed that species richness decreased quickly but not steadily along this gradient, but life-form spectra data highlighted that the high levels of plant species biodiversity in cultivated groves were due primarily to the large number of therophytes (annual plants). At a shorter time scale, we also studied the consequences of experimental clear-cutting on plant species biodiversity. Direct measurements over four years showed that species richness and diversity in the understorey increased in the first two years after clear-cutting, but decreased during the 3rd and 4th years. This decrease corresponded closely with a steady increase in Leaf Area Index of the forest canopy. These results have been used to suggest some possible ways to manage the biodiversity in these ecosystems.     

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.     

Human impact on Mediterranean wetland Coleoptera: an historical perspective at Tourves (Var,France)

Very little is known about the Neolithic human impact on the coleopteran fauna in the Mediterranean region. The site of Tourves provides an exceptional opportunity to document this major event in Provence because it offers a continuous upper Holocene sedimentary record, showing unambiguous pollen signals for anthropic activities during the Atlantic and Subboreal periods. The level dated 4310 BP is marked by a drastic change in the coleopteran assemblages: a rich marshy fauna is suddenly replaced by an impoverished fauna in which running-water elements suggest a change in the hydrological regime, probably induced by drainage of the site which may be attributed to human activities. Some wetland species such as Odacantha melanura and Cerapheles lateplagiatus, present below the level dated 4310 years BP, are now extinct in the area.     

The effect of fire on nutrient losses and cycling in aQuercus coccifera garrigue (southern France)

The effects of burning on plant nutrient budgets and rates of carbonic gas and particulate matter emission during fires were evaluated in aQuercus coccifera (garrigue) shrubland. Nutrient levels were determined in field-collected pre-fire vegetation and combustion residues. The losses (increased elemental transfer) were calculated as the difference between the quantity of an element in the fuel (combustible standing vegetation plus litter) before burning and that present in the postfire residues (ash). Weight losses of elements are correlated with weight losses of burnt plant biomass. The relative order of nutrient losses was: N>C>Na>Ca>P>K>Mg. Estimated losses of N, C and P from combustible plant matter exceeded 98, 97 and 79% respectively. Copious N, C and P volatilization during burning was promoted by high concentrations of these elements in foliage and fine woody biomass of the aboveground vegetation and leaf litter of the garrigue. Elements were principally removed in the smoke. The quantities of gaseous emissions of CO₂, CO and particulate matter produced were estimated.     

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.     

The southern limit of the Mediterranean vegetation in the Sahara during the Holocene

Holocene records from the southern Sahara in Niger allow a reconstruction of the vegetation history and inform us about the former extension of the Mediterranean. Both pollen and charcoal analyses evidenced the direct contact of Sudanian and Saharan savannas during the middle Holocene at about 19°N, whereas at 20°N the transition from the Saharan savanna to the desert was found. In southwestern Libya (26°N) a combination of a Saharan desert vegetation and a semi‐desert Artemisia shrub on the plateaus demonstrated the contact with Mediterranean influenced formations. Regular ash and charcoal layers in middle‐Holocene sediments of the northern Niger prove an early interference of man with the vegetation development. One has to imagine that, in combination with the cattle‐keeping and the later metal production, man could have changed the former northern Sudanian vegetation into the present Sahelian savanna system from the middle Holocene on.     

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.     

Impact of grazing on the species richness of plant communities in Mediterranean temporary pools (western Morocco)

The impact of grazing on the vegetation of Moroccan temporary pools has been studied at 2 scales: regional (inter-pools) and local (intra-pools). Half of the 16 forest pools studied is located in a reserve and ungrazed. The other half, located within public forest, is grazed. Vegetation relevés coupled to water-depths measurements were carried out in each pool. The results showed a significant effect of grazing on both scales of analysis. This effect was found in the species composition of the vegetation, which differed between the 2 types of pools, and in the lower species richness and abundance of plant species in the grazed pools. These differences are interpreted as resulting from the selection by herbivores and the differential tolerance of species to disturbance. These impacts are likely to expose certain species to local extinction by reducing their populations.     

Effects of afforestation with pines on woody plant diversity of Mediterranean heathlands in southern Spain

Open heathlands of the Strait of Gibraltar region constitute a singularvegetation type within the Mediterranean Basin owing to their high biodiversitylevels and distinct features in the biological aspects of endemism. However,they have been traditionally depicted as low profitable, tree-less, barrenlandsand have been either overlooked in conservation policies or included inextensive pine-tree afforestation programmes. Nowadays, most of the existingpine plantations are no longer exploited and have become dense pine woodlands.Here we present the results of a comparative analysis aimed to ascertain theeffects of pine stands on the structure and diversity of the woody component ofthese mediterranean heathlands. Ten sampling sites were located within thestudyarea, each consisting of two adjacent subsamples: an open heathland and aneighbouring heathland under pine-tree cover. Woody species richness andoverallshrub cover was much lower in pine-tree heath understoreys. This drop in coverwas more marked for short-lived species (seeders and withering resprouters),implying a decrease of the life-history diversity and also having noticeableconsequences in homogenizing the floristic composition of heath patches underpine trees. Heathland narrow endemics dropped significantly in pine stands,showing greater sensitivity to afforestation than nonendemics. We suggest thatthe open heathlands of the Strait of Gibraltar should be given furtherattentionin EU and IUCN conservation policies. Programmes should be implemented toprogressively eliminate abandoned pine plantations so as to restore andpreservethese highly diverse mediterranean heathlands.     

Copper in plant species in a copper gradient in Catalonia (North East Spain) and their potential for phytoremediation

The accumulation of Cu in roots and shoots of 32 plant species growing on soils with a wide range of Cu concentrations (30–18 500 g g^–1 total soil Cu) located in Collserola Mountain (Barcelona, Spain) was analysed. High Cu availability decreased the species diversity in the Hyparrhenietum hirto-pubescentis, the natural plant association at the study sites. Shoot and root Cu concentrations in relation to extractable soil Cu concentrations were used for the evaluation of the Cu resistance strategy in the different species. Saturation of Cu accumulation in roots was observed in most species. Hyparrhenia hirta was the most efficient shoot excluder, while the linear increase of shoot Cu with the Cu soil concentration exhibited the highest slope in Hirschfeldia incana. Most species accumulated more Cu in roots than in shoots. High shoot/root Cu ratios were only found in the highly Cu-resistant Hirschfeldia incana, in the resistant or moderately resistant Spartium junceum and Reseda sp. (R. lutea and R. phyteuma), and in the much less resistant Ononis natrix. Only two species, Hirschfeldia incana and Sedum sediforme were able to support the extreme Cu-toxicity conditions on soils with 5000–16 800 g g^–1 extractable Cu. Among the grass species tested Hyparrhenia hirta was the most Cu-resistant species (up to 1950 g g^–1 extractable soil Cu). The potential usefulness of these pseudometallophytes for phytoremediation of Cu-contaminated soils is discussed.     

Phenomorphology and ecomorphological characters of Maytenus senegalensis L. Shrublands in the Iberian Peninsula: A comparison with other Mediterranean plant communities

This work sets out to establish ecomorphological and phenomorphological patterns in the constituent flora of an Iberian Maytenus senegalensis plant community in order to relate it with or separate it from other types of Mediterranean vegetation. This plant community contains tall shrubs, scrubs, palms and vines of a variety of families. Typical ecomorphological adaptations to the Mediterranean climate were observed in the community studied: an abundance of thorny species or species with tomentose leaves and photosynthetic stems, small leaves and a high presence of species with organs capable of post-fire regeneration. However, the community also showed hybrid characteristics between scrubland (predominantly malacophyllous leaves lasting 9 months on average and low biomass) and tall shrubland communities (predominance of phanerophytes up to 3 m in height, predominantly smooth bark, periodical reposition of leaves, mean lifespan of plants 25 years and mainly fleshy fruit). The community also presents an outstanding quantity of amphiphytes (29%). From a phenological point of view, the formation of buds in winter, flowering in spring, summer–autumn seed dispersal and leaf shedding in summer are typical of other Mediterranean communities, while fructification and growth lasting three seasons, as in the case of Maytenus shrubland, is not. The RVA phenophasic index (balance between reproductive and vegetative phenophases) with values of 0.4–2 shows an intermediate value between typical shrubland (0.3–1) and forest community (1–2) values. The APC index (period of phenophasic activity of the community) differentiates it from others, since it remains at 100% for seven months, three of which are winter months. Phenophasic patterns (overlapping, or not, of flower bud formation, flowering and growth) show how similar M. senegalensis shrubland is to forests because of the absence of phenophase pattern type A (characteristic in open-scrublands); in addition, we have detected a new pattern (type G), still regarded as exclusive to M. senegalensis. As regards the existence of coenomorphs (group of species with similar morpho-functional strategies), the studied community showed much greater diversity than other communities studied to date, with an Strategies Richness index (SRi) of 0.79 (11 groups covering the 14 species existing), pointing to the co-existence of a great variety of eco-phenomorphological strategies, each with a successful response to the same ecological conditions. Finally, some of the species in the community, e.g. Asparagus horridus, Chamaerops humilis and M. senegalensis, stand out because of their unique ecomorphological and phenological characteristics.     

Effects of repeated burning on Mediterranean communities of the northeastern Iberian Peninsula

Question: In relation to a single fire, do repeated wildfires in Mediterranean type ecosystems (1) reduce plant species richness or diversity; (2) modify patterns of abundance or dominance of plant species or (3) alter plant composition? Location: Pinus halepensis dominated communities of Catalonia, northeastern Iberian Peninsula, western Mediterranean Basin. Methods: Regional, paired design with 14 study sites, each consisting of a once burnt area (1994) and a twice burnt area (1975–1993 and 1994). Ten years after the last fire, we recorded all vascular plant species present in nested plots and quantified their relative abundances on transects. We compared species richness, diversity, dominance and relative abundance and species‐area correlations between paired once and twice burnt areas and assessed their floristic composition similarity. Results: No statistically significant differences were found in species richness or diversity. Slopes of species‐area correlations were higher in once burnt areas. In twice burnt areas, dominance by one or two species was higher. P. halepensis showed lower relative abundance and nanophanerophytes showed higher relative abundance. No differences were found for resprouter, seeder or resprouter‐seeder species. Floristic composition similarity between paired areas tended to be higher in less productive sites. Conclusions: Fire recurrence had contrasting effects on species richness at different spatial scales. Repeated burning reduced the relative abundance of the dominant tree species, which resulted in a higher relative abundance of shrubs. It also promoted the dominance of herbs, particularly Brachypodium retusum. However, it did not change the relative abundance of regenerative groups. Paired areas were more similar as they were more Mediterranean in terms of climatic conditions.     

Holocene climatic changes in the Western Mediterranean, from south-east France to south-east Spain

Holocene climatic changes along coastal regions from south-east France to south-east Spain were studied using pollen ratios. Comparing modern pollen rain, vegetation and climate along selected transects from the Atlantic Ocean to the Mediterranean, we obtained threshold values of two different ratios corresponding to the different climatic conditions along the transects. These pollen ratios and threshold values were employed to characterize the Holocene climatic changes from nine Mediterranean coastal sites. The results were compared with data from marine and continental pollen sequences distributed in the western Mediterranean basin, and with additional regional data independent of human activity: lake-level fluctuations, alpine glacier advance and retreat chronology, ¹⁴C anomaly and cooling phases in Eastern France and Central Europe. The role of anthropogenic activities and climate on the changes in vegetation is discussed. Six major changes in vegetation cover were identified. They correspond to aridification phases that occurred around 9500–9000 yr BP (10 900–9700 cal BP), 7500–7000 yr BP (8400–7600 cal BP), 4500–4000 yr BP (5300–4200 cal BP), 3700–3300 yr BP (4300–3400 cal BP), 2600–1900 yr BP (2850–1730 cal BP) and 1300–1000 yr BP (1300–750 cal BP). These arid episodes were regional responses to more global climatic changes and determined the changes in the vegetation cover. Humans undoubtedly enhanced the vegetation changes, but none the less had to adapt to these new climatic conditions.     

Vulnerability of Mediterranean Basin ecosystems to climate change and invasion by exotic plant species

Aim To assess at a broad scale the vulnerability of Mediterranean vegetation to alien plant invasion under different climatic and disturbance scenarios. Location We simulated the vegetation biogeography and dynamics on five of the main islands of the Mediterranean Basin: Mallorca, Corsica, Sardinia, Crete and Lesvos. Methods We used LPJ‐GUESS, a generalized ecosystem model based on dynamic processes describing establishment, competition, mortality and ecosystem biogeochemistry. We simulated the vegetation distribution and dynamics using a set of plant functional types (PFTs) based on bioclimatic and physiological parameters, which included tree and shrub PFTs defined especially for the Mediterranean. Additionally, two invasive PFTs, an invasive tree type and an invasive herb type, were defined and used to estimate the vulnerability to invasion of a range of different ecosystems. The model was used to simulate climate changes and associated changes in atmospheric [CO₂] to 2050 according to two SpecialReport on Emissions Scenarios climate scenarios (A1Fi and B1) combined with mean disturbance intervals of 3 and 40 years. Results The simulations and scenarios showed that the effect of climate change alone is likely to be negligible in many of the simulated ecosystems, although not all. The simulated progression of an invasion was highly dependent on the initial ecosystem composition and local environmental conditions, with a particular contrast between drier and wetter parts of the Mediterranean, and between mountain and coastal areas. The rate of ecosystem disturbance was the main factor controlling susceptibility to invasion, strongly influencing vegetation development on the shorter time scale. Main conclusions Further invasion into Mediterranean island ecosystems is likely to be an increasing problem: our simulations predict that, in the longer term, almost all the ecosystems will be dominated by exotic plants irrespective of disturbance rates.     

Natural phenological variation in aspen (Populus tremula): the SwAsp collection

The genus Populus is currently the main model system for genetic, genomic, and physiological research in trees. Phenotypic variation in aspen (Populus tremula) populations growing in different environments across Sweden is expected to reflect genetic variation that is important for local adaptation. To analyze such natural phenotypic and genetic variation, the Swedish Aspen (SwAsp) Collection was established. Trees were taken from 12 different populations across Sweden, from 56° to 66° latitude north and planted in two common gardens in Ekebo (55.9°N) and Sävar (63.4°N). Data related to phenological and growth traits were collected during the second year of growth. Some traits like the date of bud set and leaf area duration showed strong clinal variation patterns with latitude in both field trials, but the date of bud flush did not change along a latitudinal cline. The phenological traits showed moderate within-populations heritabilities, although growth traits showed weaker clinal patterns and lower heritabilities than the phenological traits. This research forms the starting point for the development of the SwAsp collection, a resource facilitating analysis of the natural genetic variation in aspen, the elucidation of the structure and dynamics of aspen populations, and the future identification of the genes controlling adaptive traits using association mapping of selected candidate genes.     

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.     

Seasonal and annual variation of carbon exchange in an evergreen Mediterranean forest in southern France

We present 9 years of eddy covariance measurements made over an evergreen Mediterranean forest in southern France. The goal of this study was to quantify the different components of the carbon (C) cycle, gross primary production (GPP) and ecosystem respiration (R_eco), and to assess the effects of climatic variables on these fluxes and on the net ecosystem exchange of carbon dioxide. The Puéchabon forest acted as a net C sink of ?254 g C m^?2 yr^?1, with a GPP of 1275 g C m^?2 yr^?1 and a R_eco of 1021 g C m^?2 yr^?1. On average, 83% of the net annual C sink occurred between March and June. The effects of exceptional events such the insect‐induced partial canopy defoliation that occurred in spring 2005, and the spring droughts of 2005 and 2006 are discussed. A high interannual variability of ecosystem C fluxes during summer and autumn was observed but the resulting effect on the annual net C budget was moderate. Increased severity and/or duration of summer drought under climate change do not appear to have the potential to negatively impact the average C budget of this ecosystem. On the contrary, factors affecting ecosystem functioning (drought and/or defoliation) during March–June period may reduce dramatically the annual C balance of evergreen Mediterranean forests.     

Aeropalynological and phenological study in two different Mediterranean olive areas: Cordoba (Spain) and Perugia (Italy)

ABSTRACT

An aerobiological and phenological investigation on the olive tree was carried out during three years in two areas: Cordoba (Spain) and Perugia (Italy). In these countries, this species is economically important and those areas were chosen because of the long series of aeropalynological data (1982–1998) available, obtained by means of identical volumetric pollen traps. The aim of this study was to use phenological observations to prove the real contribution to the pollen curves in different cultivated areas. Results show that in Cordoba province (302.152 ha) the pollen curve is characterised by different peaks because of the pollination of different cultivated crops. In some cases, these crops are located far from the pollen trap (50 km) but pollen is transported thanks to favourable winds during the flowering period. In Perugia (750 ha) the pollen curve is characterised by only one peak; it is very concentrated because of the proximity of the investigated crops. The objective of this research was to obtain information on this species in order to elaborate statistical models aimed at forecasting the potential fruit production based on the amount of pollen released into the atmosphere.