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Dantas V.D.L.,Federal University of São Carlos | Pausas J.G.,CIDE CSIC
Journal of Ecology | Year: 2013

Fire and herbivory are the main disturbances shaping the structure of savannas. In these ecosystems, the key strategies by which woody plants escape fire are either early height growth (the lanky strategy) or early bark growth (the corky strategy). We hypothesize that the dominance of each strategy in different savannas depends on the prevailing disturbance regimes. Given the importance of herbivory in afrotropical savanna, we expect woody plants in these savannas to be taller and have thinner barks (the lanky strategy) than plants in neotropical savanna where fire tends to be more intense (the corky strategy). We compiled data on bark thickness and stem height in relation to stem diameter for afrotropical and neotropical savanna woody species and tested for differences in the allometric relationship between these two savannas with a general linear mixed model (GLMM). Fire intensities were higher in neotropical than in afrotropical savannas. Afrotropical savanna plants were taller and had thinner barks for a given diameter than neotropical savanna plants - supporting our hypothesis that because of the contrasting disturbance regimes, the lanky strategy is more adaptive in afrotropical savannas, whereas the corky strategy is more adaptive in neotropical savannas. Synthesis. While the lanky strategy is more associated with heavily browsed and fuel-controlled savannas, the corky strategy is associated with lightly browsed savannas that experience more intense fires. Because the relative role of disturbances varies across the globe, we suggest that the height-bark-diameter scheme is a powerful framework for understanding the ecology of many savannas. In savanna ecosystems there are two main strategies to scape fire. The lanky strategy (early alocation to height) is more associated with heavily-browsed and fuel-controlled savannas, while the corky strategy (early allocation to a thick bark) is associated with lightly-browsed savannas that experience more intense fires. Because the relative role of disturbances varies across the globe, we suggest that the height-bark-diameter scheme is a powerful framework for understanding the ecology of many savannas. © 2013 British Ecological Society.

Dantas V.d.L.,Federal University of São Carlos | Pausas J.G.,CIDE CSIC | Batalha M.A.,Federal University of São Carlos | de Paula Loiola P.,Federal University of São Carlos | Cianciaruso M.V.,Federal University of Goais
Oecologia | Year: 2013

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. © 2012 Springer-Verlag.

Pausas J.G.,CIDE CSIC | Keeley J.E.,U.S. Geological Survey | Keeley J.E.,University of California at Los Angeles
Ecosystems | Year: 2014

Wildfires have played a determining role in distribution, composition and structure of many ecosystems worldwide and climatic changes are widely considered to be a major driver of future fire regime changes. However, forecasting future climatic change induced impacts on fire regimes will require a clearer understanding of other drivers of abrupt fire regime changes. Here, we focus on evidence from different environmental and temporal settings of fire regimes changes that are not directly attributed to climatic changes. We review key cases of these abrupt fire regime changes at different spatial and temporal scales, including those directly driven (i) by fauna, (ii) by invasive plant species, and (iii) by socio-economic and policy changes. All these drivers might generate non-linear effects of landscape changes in fuel structure; that is, they generate fuel changes that can cross thresholds of landscape continuity, and thus drastically change fire activity. Although climatic changes might contribute to some of these changes, there are also many instances that are not primarily linked to climatic shifts. Understanding the mechanism driving fire regime changes should contribute to our ability to better assess future fire regimes. © 2014 Springer Science+Business Media New York.

Pausas J.G.,CIDE CSIC | Keeley J.E.,U.S. Geological Survey | Keeley J.E.,University of California at Los Angeles
New Phytologist | Year: 2014

There are two broad mechanisms by which plant populations persist under recurrent disturbances: resprouting from surviving tissues, and seedling recruitment. Species can have one of these mechanisms or both. However, a coherent framework explaining the differential evolutionary pressures driving these regeneration mechanisms is lacking. We propose a bottom-up approach in addressing this question that considers the relative survivorship of adults and juveniles in an evolutionary context, based on two assumptions. First, resprouting and seeding can be interpreted by analogy with annual versus perennial life histories; that is, if we consider disturbance cycles to be analogous to annual cycles, then resprouting species are analogous to the perennial life history with iteroparous reproduction, and obligate seeding species that survive disturbances solely through seed banks are analogous to the annual life history with semelparous reproduction. Secondly, changes in the selective regimes differentially modify the survival rates of adults and juveniles and thus the relative costs and benefits of resprouting versus seeding. Our approach provides a framework for understanding temporal and spatial variation in resprouting and seeding under crown-fire regimes. It accounts for patterns of coexistence and environmental changes that contribute to the evolution of seeding from resprouting ancestors. © 2014 New Phytologist Trust.

Verdu M.,CIDE CSIC | Pausas J.G.,CIDE CSIC
Evolution | Year: 2013

Phylogenetic methods to detect lineage diversification have been traditionally used within a particular taxonomic clade, but rarely applied to detect local diversification. For understanding in situ diversification triggered by novel conditions it is necessary to focus on the time slice where such conditions occur. These new conditions may differentially affect the diversification rate of lineages with different morpho-functional syndromes. A prominent example of these processes occurs in the Mediterranean Basin, where climate arising along the Tertiary/Quaternary transition acted as an environmental filter. In this context, lineages with different syndromes (sclerophyllous and nonsclerophyllous) are hypothesized to have different local diversification rates after the rise of the Mediterranean conditions. We used macroevolutionary methods of time-dependent diversification on a calibrated local phylogeny accommodating topological and chronological uncertainty to test syndrome-driven diversification in Mediterranean shrublands from the eastern Iberian Peninsula. We found phylogenetic evidence of higher speciation associated with the nonsclerophyllous syndrome, although extinction rates were similar between syndromes. Consequently a syndrome-driven local diversification has occurred in shrublands under Mediterranean conditions. The results provide an example of how the integration of the environmental filter in a dated phylogeny may recreate the local history of lineages and help to explain assembly processes in Mediterranean ecosystems. © 2013 The Author(s). Evolution © 2013 The Society for the Study of Evolution.

Functional Ecology | Year: 2015

Bark is a vital and very visible part of woody plants, yet only recently has bark characteristics started to be considered as key traits structuring communities and biomes. Bark thickness is very variable among woody plants, and I hypothesize that fire is a key factor selecting for a thick bark, and thus, at the global scale, a significant proportion of the variability in bark thickness is explained by the variability in fire regimes. Previous research has focused on the importance of bark thickness mainly in surface-fire regimes; here I generalize this idea and present a conceptual framework to explain how the different drivers that affect fire intensity have shaped bark thickness, in conjunction with other plant traits. I first review methods used to study bark thickness and then provide examples of bark thickness patterns from a wide range of ecosystems subject to different fire regimes (understorey fires, grass-fuelled surface fires, grass-fuelled crown fires and infrequent fires). There are some fire regimes that select for thick barks, while some only in the base of the trunk (e.g. understorey fires), others select for a thick bark on the whole plant (e.g. grass-fuelled crown fires). There are also fire regimes in which allocating resources to a thick bark is not adaptive (e.g. woody-fulled crown fires). Fire regime can explain a large proportion of the variability of bark thickness at the global scale, and thus, this trait varies across ecosystems in a predictable manner; however, the current paucity of data limits a fully accurate analysis. © 2014 The Authors.

Trends in Plant Science | Year: 2015

Microevolutionary studies show that fire traits are variable within and among populations, especially among those subject to different fire regimes. In addition, there is increasing evidence of an inherited genetic basis to variability in fire traits. Added together, pines provide compelling evidence that fire can exert an evolutionary pressure on plants and, thus, shape biodiversity. In addition, evolutionary fire ecology is providing insights to improve the management of pine forests under changing conditions. The lessons learned from pines may guide research on the evolutionary ecology of other taxa. © 2015 Elsevier Ltd.

Hernandez-Serrano A.,CIDE CSIC | Verdu M.,CIDE CSIC | Gonzalez-Martinez S.C.,CIFOR INIA | Pausas J.G.,CIDE CSIC
American Journal of Botany | Year: 2013

Premise of the study: Serotiny (delayed seed release with the consequent accumulation of a canopy seedbank) confers fitness benefits in environments with crown-fire regimes. Thus, we predicted that serotiny level should be higher in populations recurrently subjected to crown-fires than in populations where crown-fires are rare. In addition, under a high frequency of fires, space and resources are recurrently available, permitting recruitment around each mother to follow the seed rain shadow. Thus, we also predicted spatial aggregation of serotiny within populations. Methods: We compared serotiny, considering both the proportion and the age of serotinous cones, in populations living in contrasting fire regimes for two iconic Mediterranean pine species (Pinus halepensis, P. pinaster). We framed our results by quantitatively comparing the strength of the fire-serotiny relationship with previous studies worldwide. •Key results: For the two species, populations living under high crown-fire recurrence regimes had a higher serotiny level than those populations where the recurrence of crown-fires was low. For P. halepensis (the species with higher serotiny), populations in high fire recurrence regimes had higher fine-scale spatial aggregation of serotiny than those inhabiting low fire recurrence systems. The strength of the observed fire-serotiny relationship in P. halepensis is among the highest in published literature. Conclusions: Fire regime shapes serotiny level among populations, and in populations with high serotiny, recurrent fires maintain a significant spatial structure for this trait. Consequently, fire has long-term evolutionary implications at different scales, emphasizing its prominent role in shaping the ecology of pines. © 2013 Botanical Society of America.

Moreira B.,CIDE CSIC | Moreira B.,University of Coimbra | Castellanos M.C.,CIDE CSIC | Pausas J.G.,CIDE CSIC
Molecular Ecology | Year: 2014

Recurrent fires impose a strong selection pressure in many ecosystems worldwide. In such ecosystems, plant flammability is of paramount importance because it enhances population persistence, particularly in non-resprouting species. Indeed, there is evidence of phenotypic divergence of flammability under different fire regimes. Our general hypothesis is that flammability-enhancing traits are adaptive; here, we test whether they have a genetic component. To test this hypothesis, we used the postfire obligate seeder Ulex parviflorus from sites historically exposed to different fire recurrence. We associated molecular variation in potentially adaptive loci detected with a genomic scan (using AFLP markers) with individual phenotypic variability in flammability across fire regimes. We found that at least 42% of the phenotypic variation in flammability was explained by the genetic divergence in a subset of AFLP loci. In spite of generalized gene flow, the genetic variability was structured by differences in fire recurrence. Our results provide the first field evidence supporting that traits enhancing plant flammability have a genetic component and thus can be responding to natural selection driven by fire. These results highlight the importance of flammability as an adaptive trait in fire-prone ecosystems. © 2014 John Wiley & Sons Ltd.

Pausas J.G.,CIDE CSIC | Ribeiro E.,Fundacion CEAM
Global Ecology and Biogeography | Year: 2013

Aim: It has been suggested that on a global scale, fire activity changes along the productivity/aridity gradient following a humped relationship, i.e. the intermediate fire-productivity hypothesis. This relation should be driven by differing relative roles of the main fire drivers (weather and fuel) along the productivity gradient. However, the full intermediate fire-productivity model across all world ecosystems remains to be validated. Location: The entire globe, excluding Antarctica. Methods: To test the intermediate fire-productivity hypothesis, we use the world ecoregions as a spatial unit and, for each ecoregion, we compiled remotely sensed fire activity, climate, biomass and productivity information. The regression coefficient between monthly MODIS fire activity and monthly maximum temperature in each ecoregion was considered an indicator of the sensitivity of fire to high temperatures in the ecoregion. We used linear and generalized additive models to test for the linear and humped relationships. Results: Fire occurs in most ecoregions. Fire activity peaked in tropical grasslands and savannas, and significantly decreased towards the extremes of the productivity gradient. Both the sensitivity of fire to high temperatures and above-ground biomass increased monotonically with productivity. In other words, fire activity in low-productivity ecosystems is not driven by warm periods and is limited by low biomass; in contrast, in high-productivity ecosystems fire is more sensitive to high temperatures, and in these ecosystems, the available biomass for fires is high. Main conclusion: The results support the intermediate fire-productivity model on a global scale and suggest that climatic warming may affect fire activity differently depending on the productivity of the region. Fire regimes in productive regions are vulnerable to warming (drought-driven fire regime changes), while in low-productivity regions fire activity is more vulnerable to fuel changes (fuel-driven fire regime changes). © 2012 John Wiley & Sons Ltd.

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