José Raventós

Climatic trends, disturbances and short-term vegetation dynamics in a Mediterranean shrubland

Fire and erosion are two major disturbances affecting Mediterranean ecosystems. Both of them are closely related to climate. There is evidence of decreasing precipitation in the Mediterranean, particularly during summer. There are also indications of an increased variability in the rainfall distribution. Climatic changes, though show high heterogeneity at a local scale. Based on these observations, we have evaluated the following hypotheses for the Region of Valencia (East Spain). (1) During the past three decades, climatic conditions have become more favourable for wildfires and high erosivity rainfall events. We have used 30-year climate records from 97 meteorological stations to examine this. Results indicate that in general the hypothesis is true, although trends are spatially dependent. (2) The effect of high intensity rain on burned land may substantially affect short-term ecosystem composition and function, and thus successional trajectories. Based on a plot scale study, we have assessed nutrient and vegetation dynamics after burning a pyrophytic community dominated by gorse (Ulex parviflorus). Erosion following high intensity rainfall affects physicochemical soil properties. As a consequence, plant cover is reduced and specific composition affected, changing the previous relationship between obligate seeder and resprouter species.

Seasonal Dynamics of Wood Formation in Pinus Halepensis from Dry and Semi-Arid Ecosystems in Spain

The seasonal dynamics of cambial activity, wood formation, occurrence of false rings (FR), and tangential bands of resin canals (RC) were investigated in Pinus halepensis from three Mediterranean dry and semi-arid ecosystems in Spain. We collected intact tissues of phloem, cambium, and outer xylem at monthly intervals throughout 2004 from each time six trees at the three sites.Cell divisions in the cambium in all trees started before our first sampling in mid-March and ceased between November and December. Cambial activity was characterized by two maxima; one in spring and another in autumn.Trees still grew in summer but at a very low rate. The first solitary RC were formed in May or June and tangential bands of RC in June or July. In general, tangential bands of RC were observed in wider growth rings. The formation of tangential bands of RC seems to be induced by drought in the second half of the growing season. FR were formed as a result of cambial reactivation in autumn and were observed in growth rings that contained more than 50 cells in a radial row. It appears that summer drought and early autumn precipitation play an important role in false-ring formation.

Fuel characteristics and fire behaviour in mature Mediterranean gorse shrublands

Since the early 1990s, Mediterranean gorse shrublands have expanded significantly in the Mediterranean regions of Spain mainly as a result of the increase in the frequency and extension of forest fires. Mediterranean gorse (Ulex parviflorus), which has been described as a degradation stage of forest communities after fire, has also been described as a fire-prone community. Thus, its presence increases the risk that new fires might occur. In spite of this evidence, there is little information on both the composition and structural characteristics of these communities or the relationship that might exist between these vegetation characteristics and fire behaviour. In this paper we present the results of a characterization of the vegetative structure (plant density, specific composition, biomass fractions, and horizontal and vertical fuel distribution) in Mediterranean gorse. We also analyse fire behaviour using indicators obtained at different scales. Our results show mature Mediterranean gorse shrublands to be communities with high biomass values (3000–4000 g m−2) and high horizontal and vertical vegetation continuity, in which the proportion of fine dead fuel fractions with low moisture content is around 50% of the total phytomass present. Ulex parviflorus is the dominant species and its degree of dominance is a key element in the behaviour of fire. Both the fire-line intensity values and the fire severity values observed can be considered high with respect to those observed in other Mediterranean communities, thus confirming Mediterranean gorse as a high-risk community.

Size mediated climate–growth relationships in Pinus halepensis and Pinus pinea

Functional processes in trees undergo changes as the tree size increases, which may affect the response of trees to environmental factors. We tested tree-ring response to climate in four groups of trees, including large and small Pinus halepensis and Pinus pinea trees using cluster, principal component (PC) and dendroclimatological analysis. The trees were of the same age and growing on a plantation in the semiarid coastal area of southern Spain. Cluster and PC analyses showed a clear separation into four groups of trees. Autocorrelation and mean sensitivity showed significant differences between the two size classes. PCA recognised four representative principal components where PC1 represented the tree-ring—climate variability common to both species and sizes, whereas PC2, PC3 and PC4 represented a species-specific and size-dependent response of trees to climate. The differences between the two size classes were greater than those between the two species. The results suggest that future tree-ring studies should include trees stratified by size. Only this would make it possible to produce unbiased predictions on the consequences of climate change and to devise suitable mitigation strategies for preserving Mediterranean forest ecosystems.

Evidence for the spatial segregation hypothesis: a test with nine-year survivorship data in a Mediterranean shrubland

A current focus of ecology is the investigation of spatial effects on population and community dynamics; however, spatiotemporal theory remains largely untested by empirical observations or experimental studies. For example, the segregation hypothesis predicts that intraspecific aggregation should increase the importance of intraspecific competition relative to interspecific competition, thereby enhancing local coexistence in plant communities. We applied recent methods of point pattern analysis to analyze a unique long-term data set on fully mapped seedling emergence and subsequent survival in a Mediterranean gorse shrubland after experimental fires and simulated torrential rainfall events. Our overall aim was to test if the observed spatial patterns were consistent with the segregation hypothesis during the entire community dynamics from early seedling emergence to the establishment of a mature community, i.e., we explored if the observed initial segregation did indeed prevent interspecific competition from becoming dominant. We used random labeling as the null model and specific test statistics to evaluate different biological effects of the spatial interactions that determine mortality. We found that mortality was clearly not random. Comparison of the probability of mortality in dependence on the distance to conspecific and to heterospecific plants showed that mortality was controlled almost entirely by intraspecific interactions, which is consistent with the segregation hypothesis. Dead plants were aggregated and segregated from surviving plants, indicating two-sided scramble competition. Spatial interactions were density dependent and changed their sign over the course of time from positive to negative when plants grew to maturity. The simulated torrential rainfall events and subsequent erosion caused nonspecific mortality of seedlings but did not reduce the prevalence of intraspecific competition. Our results provide support for the hypothesis that the spatial distribution of plants may profoundly affect competition and can be an important determinant in the coexistence of species and biodiversity.

Fire Risk and Vegetation Structural Dynamics in Mediterranean Shrubland

Phytomass structural characteristics are highly related to vegetation flammability. In fire-prone species like Mediterranean gorse, which accumulate standing dead fuel, susceptibility to fire is a function of fuel load, vegetation composition and fuel cover, and these characteristics change with time. Thus, for effective fuel control management, knowledge of the vegetation structural dynamics related to fire risk is crucial for preventing future fires. This study analyses structural dynamics in the above-ground phytomass of Ulex parviflorus shrublands in relation to different stages of flammability, i.e., the amount of time elapsed since the last fire. For this, 152 plants were cut from shrublands at different stages of development (young, mature and senescent), and various dimensional measurements were taken on each. The phytomass was separated into living or dead fuel fractions as well as into twigs or branches depending on the stem diameter. Basal diameter is the variable that best predicted Ulex parviflorus total phytomass as well as that of the different fractions. Both dimensional and phytomass variables increased with plant development. In the young shrublands Ulex parviflorus constitutes 54% of total phytomass, and Ulex parvifloruss dead twigs fraction accounts for 5% of total phytomass. In the mature and senescent shrublands, this species represents 80% of total shrubland phytomass, and dead twigs reach values greater than 40%. Our results show that structural changes in the fuel over short periods of time (young and mature) reveal critical periods in shrub development. Identification of these stages is a necessary tool for planning fuel control programmes.

Early post-fire regeneration dynamics of Brachypodium retusum Pers. (Beauv.) in old fields of the Valencia region (eastern Spain)

Extensive wildfires have affected the Valencia region in the last two decades. A large portion of the burnt areas has been localised in old fields. Although Mediterranean communities are usually resilient to fire and recover very quickly to the pre-fire state, burnt old fields with a low number of species could show poor recovery capacity. The response capacity of these systems to fire, and, especially, the role of the native herbaceous Brachypodium retusum were studied. Two years after fire, plant cover reached a value of 56 %. Resprouter species were more important in number and in specific cover. B. retusum represented the species with the highest contribution to total plant cover for all the study period. Total biomass in burnt plots was always lower than in unburnt plots. However, B. retusum green biomass showed a quick recovery to pre-disturbance levels, and below-ground biomass was similar in burnt and unburnt plots. Reproductive biomass greatly increased in burnt with respect to unburnt plots during the first post-fire year, with no stalk production in unburnt plots. B. retusum may be considered a promising species for the restoration of extremely degraded lands because of its high resilience to fire.

Plasticity in Dendroclimatic Response across the Distribution Range of Aleppo Pine (Pinus halepensis)

We investigated the variability of the climate-growth relationship of Aleppo pine across its distribution range in the Mediterranean Basin. We constructed a network of tree-ring index chronologies from 63 sites across the region. Correlation function analysis identified the relationships of tree-ring index to climate factors for each site. We also estimated the dominant climatic gradients of the region using principal component analysis of monthly, seasonal, and annual mean temperature and total precipitation from 1,068 climatic gridpoints. Variation in ring width index was primarily related to precipitation and secondarily to temperature. However, we found that the dendroclimatic relationship depended on the position of the site along the climatic gradient. In the southern part of the distribution range, where temperature was generally higher and precipitation lower than the regional average, reduced growth was also associated with warm and dry conditions. In the northern part, where the average temperature was lower and the precipitation more abundant than the regional average, reduced growth was associated with cool conditions. Thus, our study highlights the substantial plasticity of Aleppo pine in response to different climatic conditions. These results do not resolve the source of response variability as being due to either genetic variation in provenance, to phenotypic plasticity, or a combination of factors. However, as current growth responses to inter-annual climate variability vary spatially across existing climate gradients, future climate-growth relationships will also likely be determined by differential adaptation and/or acclimation responses to spatial climatic variation. The contribution of local adaptation and/or phenotypic plasticity across populations to the persistence of species under global warming could be decisive for prediction of climate change impacts across populations. In this sense, a more complex forest dynamics modeling approach that includes the contribution of genetic variation and phenotypic plasticity can improve the reliability of the ecological inferences derived from the climate-growth relationships.

EARLY TO RISE MAKES A PLANT HEALTHY, WEALTHY, AND WISE

Seedling emergence time is a crucial event in the life cycle of a plant, determining its fitness via different components including survival, growth, and fecundity. Precocious emergents usually survive, grow, and/or reproduce earlier in the life cycle, but for perennials it is unknown whether these benefits are maintained throughout the life of the plant. Here, we examine for the first time whether fitness benefits due to the early emergence of perennial plants are perpetuated or vanish with time. For nine years, in a Mediterranean gorse community, we followed the fate of 2118 seedlings belonging to the four dominant woody species. We estimated phenotypic selection gradients on emergence time for three fitness components (survival, growth, and fecundity), under two experimentally simulated scenarios (fire and fire + erosion), at two different times in the life of the plant (3 and 9 years). Fire and erosion represent two potential selective forces constraining the temporal window of seedling emergence in Mediterranean habitats. All the species exhibited selection for early emergence, but through different fitness components. Directional selection favoring early emergence via survival in both fire scenarios was detected in the two Cistaceae species (Helianthemum marifolium and Cistus albidus), in which precocious emergents had higher fitness values late in the life cycle (9 years). In contrast, Fabaceae species (Ononis fruticosa and Ulex parviflorus) were not selected for early emergence via survival. Early emergents of all species in both fire scenarios had higher fitness values through growth early in the life cycle; these benefits decreased slightly with time but remained statistically significant, except in H. marifolium. Finally, late fecundity was enhanced by early emergence in both fire scenarios in C. albidus and U. parviflorus but not in H. marifolium. In conclusion, benefits acquired by emerging early are perpetuated for at least nine years.

Fire and torrential rainfall: effects on the perennial grass Brachypodium retusum

Summer wildfires and autumn torrential rainfall are the background to the development of one of the most important environmental problems in the western Mediterranean area: erosion, degradation of soil structure and desertification. These processes especially affect plant communities located in old abandoned fields where woody sprouters are practically absent. Under these conditions, post-fire vegetation cover is dependent on obligate seeders and, in the short term, especially on resprouting perennial grasses. Brachypodium retusum is a rhizomatous perennial grass that plays a particularly important role in the resilience of these fire-prone Mediterranean ecosystems. In Mediterranean gorse shrublands, during the first few years after a fire, this species represents 90% of plant cover and biomass. The recurrence or severity of fire does not seem to affect the regeneration capacity of this species. After two years, even under high fire frequency (12 years) and high fire severity (> 400 °C), it recovers both cover and biomass. Nevertheless, this species, a key factor in the control of soil erosion, is extremely affected by erosion processes. A single extreme rainfall event reduces the rate of regeneration of B. retusum by about 50%. The reduction in the protective cover of this species may result in degradation and eventual self-induced degrading process in the ecosystem.