Susana Paula

Fire‐related traits for plant species of the Mediterranean Basin

Plant trait information is essential for understanding plant evolution, vegetation dynamics, and vegetation responses to disturbance and management. Furthermore, in Mediterranean ecosystems, changes in fire regime may be more relevant than direct changes in climatic conditions, making the knowledge of fire-related traits especially important. Thus the purpose of this data set was to compile the most updated and comprehensive information on fire-related traits for vascular plant species of the Mediterranean Basin, that is, traits related to plant persistence and regeneration after fire. Data were collected from an extensive literature review and from field and experimental observations. The data source is documented for each value. Since life history traits may vary spatially or with environmental conditions, we did not aggregate them by species; i.e., traits and species are repeated in different records if they were observed by different researchers and/or in different locations. Life history traits inclu...

Fuel loading and flammability in the Mediterranean Basin woody species with different post-fire regenerative strategies

The flammability and combustibility of plant communities are determined by species features related to growth-form, structure and physiology. In some ecosystems, such as the Mediterranean ones, these characteristics may contribute to the existence of fire-prone species. We measured several parameters associated with the flammability and fuel loading of dominant woody species with different post-fire regenerative strategies (seeders and non-seeders) in shrublandsinthewesternMediterraneanBasin.Overall,seederspeciesshowlowerfuelloadbutaremorepronetoburning owing to ahigher dead-to-live fuelratio, live fine-fuel proportionand dead fine-fuel proportion.Moreover, theyburst into flame at lower temperatures than non-seeders. In the Mediterranean Basin, most seeder species emerged mainly during the Quaternary, under a highly fluctuating Mediterranean climate and during recurrent fires. We propose that properties related to the combustibility and flammability of seeders may be the result of selective pressures associated with both fire andclimate.Theseresultssuggestthatecosystemsdominatedbyseederspeciesaremoresusceptibletofireriskthanthose dominated by non-seeder species in the Mediterranean Basin. Therefore, the proportion of these types of species resulting from previous fire or management history is likely to determine the characteristics of future fire events.

Towards understanding resprouting at the global scale.

Understanding and predicting plant response to disturbance is of paramount importance in our changing world. Resprouting ability is often considered a simple qualitative trait and used in many ecological studies. Our aim is to show some of the complexities of resprouting while highlighting cautions that need be taken in using resprouting ability to predict vegetation responses across disturbance types and biomes. There are marked differences in resprouting depending on the disturbance type, and fire is often the most severe disturbance because it includes both defoliation and lethal temperatures. In the Mediterranean biome, there are differences in functional strategies to cope with water deficit between resprouters (dehydration avoiders) and nonresprouters (dehydration tolerators); however, there is little research to unambiguously extrapolate these results to other biomes. Furthermore, predictions of vegetation responses to changes in disturbance regimes require consideration not only of resprouting, but also other relevant traits (e.g. seeding, bark thickness) and the different correlations among traits observed in different biomes; models lacking these details would behave poorly at the global scale. Overall, the lessons learned from a given disturbance regime and biome (e.g. crown-fire Mediterranean ecosystems) can guide research in other ecosystems but should not be extrapolated at the global scale.

Unearthing belowground bud banks in fire‐prone ecosystems

Despite long-time awareness of the importance of the location of buds in plant biology, research on belowground bud banks has been scant. Terms such as lignotuber, xylopodium and sobole, all referring to belowground bud-bearing structures, are used inconsistently in the literature. Because soil efficiently insulates meristems from the heat of fire, concealing buds below ground provides fitness benefits in fire-prone ecosystems. Thus, in these ecosystems, there is a remarkable diversity of bud-bearing structures. There are at least six locations where belowground buds are stored: roots, root crown, rhizomes, woody burls, fleshy swellings and belowground caudexes. These support many morphologically distinct organs. Given their history and function, these organs may be divided into three groups: those that originated in the early history of plants and that currently are widespread (bud-bearing roots and root crowns); those that also originated early and have spread mainly among ferns and monocots (nonwoody rhizomes and a wide range of fleshy underground swellings); and those that originated later in history and are strictly tied to fire-prone ecosystems (woody rhizomes, lignotubers and xylopodia). Recognizing the diversity of belowground bud banks is the starting point for understanding the many evolutionary pathways available for responding to severe recurrent disturbances.

Forests and Global Change: Global change and Mediterranean forests: current impacts and potential responses

Global change exacerbating Mediterranean stresses Mediterranean forests have always had to cope with challenging environmental conditions that change across different temporal and spatial scales. However, the rapidity of current environmental change, driven by greater-than-ever human influences on natural processes, is unprecedented and has triggered renewed research endeavour into the impacts on Mediterranean ecosystems (Valladares 2008). The climate of Mediterranean areas is expected to become drier and warmer, with decreasing water availability for plants and increasing evapotranspiration (IPCC 2007). This will result in more acute physiological stress, increased importance of species-specific tolerances, plasticity and thresholds, phenological change and recruitment effects (Montserrat-Martin et al . 2009; Morin et al . 2010; Penuelas et al . 2004). Several studies have demonstrated how the conditions currently experienced by seedlings and saplings are quite different to those when current adults recruited (Lloret & Siscart 1995; Montoya 1995). The anticipated impacts of such changes have led to a renewed interest in classic ecophysiological research into drought stress and tolerance (Wikelskia & Cooke 2006), as well as population-level studies on phenotypic plasticity and the evolution of tolerance in certain key tree species, such as Holm ( Quercus ilex ) and cork oaks ( Q. suber ) (Gimeno et al . 2009; Ramirez-Valiente et al . 2010). Niche modelling techniques are used to forecast changes to species distributions under future climate scenarios, and the results predict abrupt shifts of dominant tree species in the next decades. Forest diebacks, species migration and displacement, and altitudinal shifts of forest types have already been recorded (Penuelas & Boada 2003; Allen et al . 2010). For example, in northeast Spain Fagus sylvatica and Calluna vulgaris are being replaced by Quercus ilex at high elevations (Penuelas & Boada 2003).

Effects of Climate and Extreme Events on Wildfire Regime and Their Ecological Impacts

Fire regime has been affected by climate changes in the past, and is expected to do so in relation to the projected climate warming in the near future. For the Mediterranean Basin, higher fire risk, longer fire season, and more frequent large, severe fires are expected. The projected increased drought for the Mediterranean Basin would make ecosystems more vulnerable to fire, and more difficult to restore after fire. Ecosystem vulnerability is assessed considering soil susceptibility to post-fire erosion, and vegetation capacity to recover after fire.

Lignotubers in Mediterranean basin plants

Lignotubers are swollen woody structures located at the root-shoot transition zone and contain numerous dormant buds and starch reserves. This structure enables the plant to resprout prolifically after severe disturbances that remove the aboveground biomass. These are considered adaptive traits in ecosystems with highly frequent and severe disturbances—such as fire-prone ecosystems. In this paper, we aim to contribute to the knowledge of lignotubers in the Mediterranean basin and highlight the evolutionary implications. We first summarise existing knowledge on lignotuber species in the Mediterranean basin. We then provide a detailed morpho-anatomical description of early lignotubers in two common woody species (Arbutus unedo L. and Phillyrea angustifolia L.). Finally, we compare our anatomical results with those obtained in studies conducted with other lignotuberous species from different Mediterranean regions. Lignotubers were verified in 14 species in the Mediterranean basin; all being from lineages with origins dating to the Tertiary and thus pre-dating the Mediterranean climate. In A. unedo and P. angustifolia, lignotubers are macroscopically discernible in 4- and 2-year-old saplings, respectively. In these two species, the lignotubers have numerous buds protected by hypertrophied scales, and have a contorted xylem containing abundant starch. Our results challenge the traditional idea that pre-Mediterranean lineages suffered evolutionary inertia; instead, lignotuberous species may be considered examples of plants that adapted to the increased fire activity that occurred throughout the Tertiary and Quaternary. We also highlight the use of morpho-anatomical traits to unambiguously distinguish between lignotuberous and non-lignotuberous resprouting species.

Enhancement of photosynthesis in post-disturbance resprouts of two co-occurring Mediterranean Erica species

The higher growth rates of resprouting shoots compared with those of mature plants in resprouter woody species are supported by higher rates of photosynthesis and transpiration. In this contribution we hypothesize that species with higher resprouting vigour will show a larger enhancement of photosynthesis in resprouting shoots. We test this hypothesis by comparing gas exchange and leaf parameters between resprouting and mature plants in Erica scoparia and E. australis. These two Erica species co-occur in Mediterranean heathlands of the Strait of Gibraltar. Erica scoparia has a higher rate of post-disturbance starch recovery than E. australis, which makes it more resistant to recurrent disturbance. We tested the hypothesis that enhancement of photosynthesis and water use characteristics of resprouting shoots compared with mature plants should be more pronounced in E. scoparia. In both species, resprouts had higher efficiency in the use of light and higher maximum net photosynthesis than mature shoots. However, contrary to expectations, differences in the photosynthetic performance between resprouts and mature plant shoots were larger in E. australis. Higher root to shoot ratios in resprouting E. australis plants, determined by their slower above-ground recovery, together with stronger demand from carbon sinks might explain this result.

Light Environment Has Little Effect on Heteroblastic Development of the Temperate Rain Forest Tree Gevuina avellana Mol. (Proteaceae)

Premise of research. Heteroblastic species are those that show an abrupt change in shape and/or size among individual metamers during ontogeny. Gevuina avellana Mol. (Proteaceae) is a typical tree species in the temperate rain forests of Chile and Argentina. This tree shows drastic heteroblasty, changing from simple leaves at the seedling stage to pinnate leaves during development. It regenerates mostly in shady understories, but juveniles can be found growing under a wide range of light conditions (5%–50% canopy openness). Thus, considering that light has been proposed as a driver of the heteroblastic strategy, G. avellana is an interesting model to study the potential environmental modulation of its ontogenetically programmed heteroblasty. Therefore, the aim of this study was to determine the effect of light availability on G. avellana’s heteroblastic trajectory. We postulated that G. avellana’s ontogenetic changes in leaf complexity (i.e., heteroblasty) increase under high light availability. Methodology. Saplings along most of the light availability gradient were sampled. Plant height was used as a proxy for ontogeny. We measured several leaf traits (leaf area), shape (aspect ratio), pinnation (leaf dissection index), and complexity (fractal dimension index). First, we evaluated the change in each leaf trait with height by means of Pearson’s correlation. Then we tested for differences in leaf traits along the ontogeny between two light environments (higher and lower than 10% canopy openness) by the line-fitting standardized major axis method. Pivotal results. We found positive correlations between each leaf trait and plant height (P < 0.001), while only leaf size and complexity showed higher ontogenetic increases under high light. Conclusions. Light environment has a small but significant effect on the heteroblastic trajectory of G. avellana. In particular, saplings inhabiting microsites with high light availability have larger and more complex leaves. While allometry would explain the ontogenetic trajectory of leaf size, the increased complexity could reflect functional advantages for large leaves in tall plants, especially under high light conditions.

Postfire responses of the woody flora of Central Chile: Insights from a germination experiment

Fire is a selective agent shaping plant traits and community assembly in fire-prone ecosystems. However, in ecosystems with no fire history, it can be a cause of land degradation when it is suddenly introduced by humans, as plant species may not be able to respond to such novel disturbance. Unlike other Mediterranean-type ecosystems (MTE) of the world, natural fires have not been frequent during the Quaternary in the matorral of Central Chile, and thus, plant adaptive responses are expected to be uncommon. We evaluated the effect of heat shock on seed survival and germination of 21 native woody plants of the Chilean matorral and compiled information on smoke-stimulation and resprouting, to evaluate the importance of fire-adaptive responses in the context of the other MTE. We found that in the Chilean woody flora negative seed responses to fire cues were more frequent than positive responses. Although resprouting is a relatively widespread trait, fire-stimulated germination is not as common in the Chilean matorral as in other MTE. The seeds of seven endemic species were strongly damaged by fire cues and this should be considered in post-fire restoration planning. However, our results also showed that many species were resistant to elevated doses of heat shock and in some, germination was even stimulated. Thus, future research should focus on the evolutionary causes of these responses. These findings could help to develop strategies for fire management in the Chilean matorral. In addition, they will improve our understanding of the evolutionary forces that shaped this plant community and to better frame this region among the other MTE worldwide.