Adaptive flammability syndromes in thermo-Mediterranean vegetation, captured by alternative resource-use strategies.

Fire affects and is affected by leaf functional traits indicative of resource allocation trade-offs. Global change drivers constrain both the resource-use strategies and flammability of coexisting species. However, small attention has been given in identifying links among flammability and plant economics. Ambiguity comes from the fact that flammability is a multidimensional trait. Different flammability attributes (i.e. ignitibility, sustainability, combustibility and consumability) have been used to classify species, but no widely-accepted relationships exist between attributes. We hypothesised that flammability is a spectrum (defined by its four attributes) and the alternative flammability syndromes of coexisting species can be captured by their resource-use strategies. Furthermore, we argue that flammability syndromes are adaptive strategies that ensure persistence in the post-fire community. We conducted a large-scale study to estimate all flammability attributes on leaves from nine, dominant, thermo-Mediterranean species with alternative resource-use and fire-response strategies across a wide environmental and geographic gradient. We assessed the interdependence among attributes, and their variation across ecological scales (genus, species, individual, site and region). Furthermore, we collected 10 leaf functional traits, conducted a soil study and extracted long-term climatological data to quantify their effect on flammability attributes. We found that leaf flammability in thermo-Mediterranean vegetation is a continuous two-dimensional spectrum. The first dimension, driven by leaf shape and size, represents heat release rate (combustibility vs. sustainability), while the second, controlled by leaf economics, presents ignition delay and total heat release (i.e. consumability). Alternative flammability syndromes can increase fitness in fire-prone communities by offering qualitative differences in survival or reproduction. Trade-offs and constraints that control the distribution of resource-use strategies across environmental gradients appeared to drive leaf flammability syndromes as well. Tying the flammability spectrum with resource allocation trade-offs on a global scale can help us predict future ecosystem properties and fire regimes and illustrate evolutionary constraints on flammability.

Functional Trait Variation Among and Within Species and Plant Functional Types in Mountainous Mediterranean Forests.

Plant structural and biochemical traits are frequently used to characterise the life history of plants. Although some common patterns of trait covariation have been identified, recent studies suggest these patterns of covariation may differ with growing location and/or plant functional type (PFT). Mediterranean forest tree/shrub species are often divided into three PFTs based on their leaf habit and form, being classified as either needleleaf evergreen (Ne), broadleaf evergreen (Be), or broadleaf deciduous (Bd). Working across 61 mountainous Mediterranean forest sites of contrasting climate and soil type, we sampled and analysed 626 individuals in order to evaluate differences in key foliage trait covariation as modulated by growing conditions both within and between the Ne, Be, and Bd functional types. We found significant differences between PFTs for most traits. When considered across PFTs and by ignoring intraspecific variation, three independent functional dimensions supporting the Leaf-Height-Seed framework were identified. Some traits illustrated a common scaling relationship across and within PFTs, but others scaled differently when considered across PFTs or even within PFTs. For most traits much of the observed variation was attributable to PFT identity and not to growing location, although for some traits there was a strong environmental component and considerable intraspecific and residual variation. Nevertheless, environmental conditions as related to water availability during the dry season and to a smaller extend to soil nutrient status and soil texture, clearly influenced trait values. When compared across species, about half of the trait-environment relationships were species-specific. Our study highlights the importance of the ecological scale within which trait covariation is considered and suggests that at regional to local scales, common trait-by-trait scaling relationships should be treated with caution. PFT definitions by themselves can potentially be an important predictor variable when inferring one trait from another. These findings have important implications for local scale dynamic vegetation models.

An integrated phenotypic trait-network in thermo-Mediterranean vegetation describing alternative, coexisting resource-use strategies.

Vascular plants have been found to align along globally-recognised resource-allocation trade-offs among specific functional traits. Genetic constrains and environmental pressures limit the spectrum of viable resource-use strategies employed by plant species. While conspecific plants have often been described as identical, intraspecific variation facilitates species coexistence and evolutionary potential. This study attempts to link an individual's phenotype to its environmental tolerance and ecosystem function. We hypothesised that: (1) seasonal variation in water availability has selected for tight phenotypic integration patterns that shape Mediterranean vegetation; however, (2) coexisting species employ alternative resource-use strategies to avoid competitive exclusion; specifically (3) species with smaller climatic niches (i.e. potential distributions) display higher functional diversity. We examined the interdependence among and the sources of variation within 11 functional traits, reflecting whole-plant economics (e.g. construction costs, hydraulics, defences, water storage capacity), from nine dominant, thermo-Mediterranean species measured across a wide environmental and geographic gradient. Furthermore, we delineated the phenotypic and climatic hypervolumes of each studied species to test for climatic niche overlap and functional distinctiveness. By adopting this multidimensional trait-based approach we detected fundamental phenotypic integration patterns that define thermo-Mediterranean species regardless of life history strategy. The studied traits emerged intercorrelated shaping a resource-allocation spectrum. Significant intraspecific variability in most measured traits allowed for functional distinctiveness among the measured species. Higher functional diversity was observed in species restricted within narrower climatic niches. Our results support our initial hypotheses. The studied functional traits collectively formed an integrated space of viable phenotypic expressions; however, phenotypic plasticity enables functionally distinctive species to succeed complementary in a given set of environmental conditions. Functional variability among coexisting individuals defined species' climatic niches within the trait-spectrum permitted by Mediterranean conditions. Ultimately, a species establishment in a locality depends on the extent that it can shift its trait values.