Disentangling plasticity of serotiny, a key adaptive trait in a Mediterranean conifer.

PREMISE OF THE STUDY: Serotiny, the maintenance of ripe seeds in closed fruits or cones until fire causes dehiscence, is a key adaptive trait of plants in fire-prone ecosystems, but knowledge of phenotypic plasticity for cone retention in woody plants is extremely scarce. On the basis of published literature and our field observations, we hypothesized that increased aridity might decrease the aerial seed bank as a plastic response, not necessarily adaptive. METHODS: We used a Pinus halepensis common garden replicated in three contrasted sites (mild, cold, and dry) to separate population differentiation from phenotypic plasticity of cone serotiny and canopy cone bank (CCB). Differences in growth among trees of the same provenance allowed us to include size effect as a proxy of ontogenetic age for the same chronological age of the trees. KEY RESULTS: Tree size had a strong negative effect on serotiny, but serotiny degree differed among trial sites even after accounting for size effects. As hypothesized, serotiny was lower at the harsh (dry and cold) sites compared with the mild site. Genetic variation for size-dependent cone serotiny and significant population × site interaction were confirmed, the latter implying different plasticity of serotiny among populations. Population differentiation for CCB showed an ecotypic trend, with positive correlation with temperature oscillation (continentality) and negative correlation with summer rainfall. CONCLUSIONS: Growth-limiting environments exacerbated the precocious release of seeds, contrary to the ecotypic trend found for the aerial cone bank, suggesting a counter-gradient plasticity. This plastic response is potentially maladaptive under a scenario of frequent wildfires.

How Does Water Availability Affect the Allocation to Bark in a Mediterranean Conifer?

Bark thickness is a key structural feature in woody plants in the protection against fire. We used 19 provenances of Pinus halepensis, an obligate-seeder species, in a replicated common garden at two environments contrasting in water availability to assess the interacting effects of site environment and population in the relative allocation to bark, expecting lower allocation at the drier site. Secondly, given the average fire frequency, we analyzed whether trees reached the critical absolute thickness soon enough for population persistence via aerial seed bank. Our analyses indicated that trees at the moister site allocated a rather fixed quantity of resources independent of tree size, and almost all populations reached critical absolute bark thickness to eventually survive fire. In contrast, at the drier site allocation to bark reduced with tree size, and most populations did not reach the critical bark thickness. Populations from areas with higher fire frequency had thicker basal bark, while those from areas with severe droughts and short vegetative periods, had thinner bark. In conclusion, drought-stressed trees have a higher risk to die from fires before achieving reproduction and building a sufficient aerial seed bank.

Maintenance costs of serotiny in a variably serotinous pine: The role of water supply.

Serotiny is an important adaptation for plants in fire-prone environments. However, different mechanisms also induce the opening of serotinous cones in the absence of fire in variably serotinous species. Xeriscence -cone opening driven by dry and hot conditions- is considered to be mediated only by the external environment, but endogenous factors could also play a significant role. Using the variably serotinous Pinus halepensis as our model species, we determined the effects of cone age and scales density in cone opening, and using in-situ and ex-situ manipulative experiments we investigated the role of water availability in the opening of serotinous cones. We hypothesized that loss of connection between the cones and the branch through the peduncles or the absence of water supply could induce a faster cone opening. Results showed that older cones lost more water and opened at lower temperatures, with no influence of scales density. Both field and chamber manipulative experiments (using paired cones of the same whorl) confirmed that water intake through the peduncles affected significantly the pace of cone opening, such that lack of water supply speeded up cone dehiscence. However, this was true for weakly serotinous provenances-more common in this species-, while highly serotinous provenances were indifferent to this effect in the field test. All our results support that cone serotiny in P. halepensis involves the allocation of water to the cones, which is highly consistent with the previously observed environmental effects. Importantly, the existence of maintenance costs of serotinous cones has strong implications on the effects of climate change in the resilience of natural populations, via modifications of the canopy seed banks and recruitment after stand-replacing fires. Moreover, evolutionary models for serotiny in P. halepensis must take into account the significant contribution of maintenance costs to the complex interaction between genotype and the environment.