The role of fire in structuring trait variability in Neotropical savannas

Intraspecific trait variability plays a fundamental role in community structure and dynamics; however, few studies have evaluated its relative importance to the overall response of communities to environmental pressures. Since fire is considered a key factor in Neotropical savannas, we investigated to what extent the functional effects of fire in a Brazilian savanna occurs via intra- or interspecific trait variability. We sampled 12 traits in communities subjected to three fire regimes in the last 12 years: annual, biennial, and protected. To evaluate fire’s relative effects, we fitted a general linear mixed models with species as random and fire as fixed factors, using: (1) all species in the communities (i.e., considering intra- and interspecific variabilities); (2) 18 species common to all fire regimes (i.e., intraspecific variability only); and (3) all species with their overall average trait values (i.e., interspecific variability only). We assessed the relative role of intra- or interspecific variability by comparing the significance of each trait in the three analyses. We also compared the within and between fire variabilities with a variance component analysis. Five traits presented larger intraspecific than interspecific variability, and the main effect of fire occurred at the intraspecific level. These results confirm that it is important to consider intraspecific variability to fully understand fire-prone communities. Moreover, trait variability was larger within than among fire regimes. Thus, fire may act more as an external filter, preventing some of the species from the regional pool from colonizing the cerrado, than as an internal factor structuring the already filtered cerrado communities.     

Fire as a key driver of Earth's biodiversity

Many terrestrial ecosystems are fire prone, such that their composition and structure are largely due to their fire regime. Regions subject to regular fire have exceptionally high levels of species richness and endemism, and fire has been proposed as a major driver of their diversity, within the context of climate, resource availability and environmental heterogeneity. However, current fire‐management practices rarely take into account the ecological and evolutionary roles of fire in maintaining biodiversity. Here, we focus on the mechanisms that enable fire to act as a major ecological and evolutionary force that promotes and maintains biodiversity over numerous spatiotemporal scales. From an ecological perspective, the vegetation, topography and local weather conditions during a fire generate a landscape with spatial and temporal variation in fire‐related patches (pyrodiversity), and these produce the biotic and environmental heterogeneity that drives biodiversity across local and regional scales. There have been few empirical tests of the proposition that ‘pyrodiversity begets biodiversity’ but we show that biodiversity should peak at moderately high levels of pyrodiversity. Overall species richness is greatest immediately after fire and declines monotonically over time, with postfire successional pathways dictated by animal habitat preferences and varying lifespans among resident plants. Theory and data support the ‘intermediate disturbance hypothesis’ when mean patch species diversity is correlated with mean fire intervals. Postfire persistence, recruitment and immigration allow species with different life histories to coexist. From an evolutionary perspective, fire drives population turnover and diversification by promoting a wide range of adaptive responses to particular fire regimes. Among 39 comparisons, the number of species in 26 fire‐prone lineages is much higher than that in their non‐fire‐prone sister lineages. Fire and its byproducts may have direct mutagenic effects, producing novel genotypes that can lead to trait innovation and even speciation. A paradigm shift aimed at restoring biodiversity‐maintaining fire regimes across broad landscapes is required among the fire research and management communities. This will require ecologists and other professionals to spread the burgeoning fire‐science knowledge beyond scientific publications to the broader public, politicians and media.     

Pines and oaks in the restoration of Mediterranean landscapes of Spain: New perspectives for an old practice — a review

Pines have been extensively used for land restoration in the Mediterranean basin and in other parts of the world, since the late 19^th century. The theoretical basis supporting pine utilisation was its stress-tolerant and pioneer features, and their attributed role of facilitating the development of late-successional hardwoods in the long-term. In the present work, the use of pines and hardwoods in forest restoration is discussed in the frame of the current disturbance regime and social demands for Mediterranean forests. Large pine plantations have recently disappeared because of their sensitivity to fire (e.g., Pinus nigra) or because of the short fire-intervals (e.g., Pinus halepensis). Combined pine and oak plantations are proposed for degraded land restoration on the basis of the complementary features of both groups of species. Seeding and containerised seedling plantation, soil amendments and plantation techniques to reduce transplant shock are evaluated for reforestation under water-stressing conditions, on the basis of several experiments performed in eastern Spain. Both P. halepensis and Quercus ilex are tested.     

Acorn dispersal estimated by radio-tracking

Bird-dispersed seeds are difficult to track, especially in the case of long-distance dispersal events. To estimate the oak dispersal distance and the seed shadow generated by the European jay (Garrulus glandarius), we inserted radio-transmitters in 239 acorns, placed them in bird-feeders and then located them by radio-tracking. Using this methodology we located the exact caching site of 94 Quercus ilex and 54 Q. suber acorns and determined the caching habitat characteristics (vegetation type, distance, spatial distribution). The results show that: (1) there is no differences in the dispersal distance distribution between the different acorn species or sizes, (2) dispersal distances range from approximately 3 m up to approximately 550 m (mean = 68.6 m; median = 49.2 m), (3) recently abandoned fields and forest tracks were the sites preferred by jays to cache acorns, whereas fields and shrublands were avoided and (4) seed shadows showed acorn aggregation zones (i.e. clusters of caches) close to the feeder as well as isolated caches at longer distances. The results also suggest that radio-transmitters are a cheap and reliable way to determine seed shadows and quantify both seed dispersal and post-dispersal seed predation for medium to large seeds.     

Scale-dependent segregation of seeders and resprouters in cork oak (<Emphasis Type="Italic">Quercus suber</Emphasis>) forests

Recent studies showed that disturbances and water availability determine the richness among plants with different post-fire strategies of Mediterranean-type ecosystems. The aim of this study was to determine whether or not the scale of analysis has an influence on the effects of these factors and, therefore, on the segregation of the dominant post-fire strategies, obligate seeders and obligate resprouters, and facultative species. We recorded all woody species and geographical features on 94 (75 m²) plots of cork oak woodlands in the southern Iberian Peninsula. For each regenerative type (resprouters, seeders and species with both traits—facultative species), we tested the relationship between the number of species and the predictors using a generalised linear mixed model. The fixed predictor considered at the large scale was altitude, and fixed predictors considered at the local scale were aspect (north/south) and disturbance (fire and clearing by heavy machinery; yes/no). The random predictor was the factor of site. When this factor did not have significant effect for some regenerative types, these relationships was tested using a generalised linear model. Resprouting species were most represented at lower altitudes and in undisturbed sites, while seeders were also at lower altitudes but mostly on south-facing slopes, especially south-facing disturbed sites. For facultative species, site is the most important variable. The proportion of seeders from the total species is not related to altitude, but it is related to disturbance and aspect. These results suggest that there is no segregation of the richness of seeders and resprouters at the large scale (altitudinal gradient). Differences appeared at the local scale (aspect and disturbance).     

Fire-adapted traits of Pinus arose in the fiery Cretaceous

? The mapping of functional traits onto chronograms is an emerging approach for the identification of how agents of natural selection have shaped the evolution of organisms. Recent research has reported fire-dependent traits appearing among flowering plants from 60 million yr ago (Ma). Although there are many records of fossil charcoal in the Cretaceous (65-145 Ma), evidence of fire-dependent traits evolving in that period is lacking. ? We link the evolutionary trajectories for five fire-adapted traits in Pinaceae with paleoatmospheric conditions over the last 250 million yr to determine the time at which fire originated as a selective force in trait evolution among seed plants. ? Fire-protective thick bark originated in Pinus c. 126 Ma in association with low-intensity surface fires. More intense crown fires emerged c. 89 Ma coincident with thicker bark and branch shedding, or serotiny with branch retention as an alternative strategy. These innovations appeared at the same time as the Earth's paleoatmosphere experienced elevated oxygen levels that led to high burn probabilities during the mid-Cretaceous. ? The fiery environments of the Cretaceous strongly influenced trait evolution in Pinus. Our evidence for a strong correlation between the evolution of fire-response strategies and changes in fire regime 90-125 Ma greatly backdates the key role that fire has played in the evolution of seed plants.     

Fire severity and seedling establishment in Pinus halepensis woodlands, eastern Iberian Peninsula

Given the observed heterogeneity in fire severity produced within wildfires, we asked to what extent this heterogeneity might affect post-fire regeneration. For this purpose, we studied the post-fire dynamics of Pinus halepensis (Aleppo pine) in the eastern Iberian Peninsula. Sampling was stratified on the basis of fire severity. We defined three fire severity classes based on the degree of consumption of the pine canopy. The results suggested that there is no clear relationship between seedling density and fire severity; however, mortality was lower and growth (height, shoot biomass and root biomass) was higher in the high severity class. These results can be explained by soil processes: Sites in the high fire severity class may have sustained higher fire intensities, resulting in higher soil organic matter mineralisation and higher ash deposition, and thus in higher post-fire soil fertility. This higher fertility would produce faster growth in pine seedlings. Independent of the severity class, seedling mortality was higher in quadrats (50 × 50 cm) with higher cover of the perennial grass Brachypodium retusum (Poaceae), suggesting a possible competitive effect. For all plots in all 3 severity classes, spatial analysis suggests an aggregate seedlings pattern, but with independence from the position of the adult (source) trees. This revised version was published online in June 2006 with corrections to the Cover Date.