Above- and below-ground trait coordination in tree seedlings depend on the most limiting resource: a test comparing a wet and a dry tropical forest in Mexico.

The study of above- and below-ground organ plant coordination is crucial for understanding the biophysical constraints and trade-offs involved in species' performance under different environmental conditions. Environmental stress is expected to increase constraints on species trait combinations, resulting in stronger coordination among the organs involved in the acquisition and processing of the most limiting resource. To test this hypothesis, we compared the coordination of trait combinations in 94 tree seedling species from two tropical forest systems in Mexico: dry and moist. In general, we expected that the water limitation experienced by dry forest species would result in stronger leaf-stem-root coordination than light limitation experienced by moist forest species. Using multiple correlations analyses and tools derived from network theory, we found similar functional trait coordination between forests. However, the most important traits differed between the forest types. While in the dry forest the most central traits were all related to water storage (leaf and stem water content and root thickness), in the moist forest they were related to the capacity to store water in leaves (leaf water content), root efficiency to capture resources (specific root length), and stem toughness (wood density). Our findings indicate that there is a shift in the relative importance of mechanisms to face the most limiting resource in contrasting tropical forests.

Differentiation in the water-use strategies among oak species from central Mexico.

Oak species (Fagaceae: Quercus) differ in their distribution at the landscape scale, specializing to a certain portion of environmental gradients. This suggests that functional differentiation favors habitat partitioning among closely related species. To elucidate the mechanisms of species coexistence in oak forests, we explored patterns of interspecific variation in functional traits involved in water-use strategies. We tested the hypothesis that oak species segregate along key trade-offs between xylem hydraulic efficiency and safety, and between hydraulic safety and drought avoidance capacity, leading to species niche partitioning across a gradient of aridity. To do so, we quantified biophysical and physiological traits in four red and five white oak species (sections Lobatae and Quercus, respectively) across an aridity gradient in central Mexico. We also explored the trade-offs guiding species differentiation, particularly between the drought tolerance versus water acquisition capacity, and determined whether the water-use strategy was associated with the portion of the environmental gradient that the species occupy. In a trait-by-trait analysis, we detected differences between white and red oak species. However, a larger part of the variation was explained at the species rather than at the section level. We detected two primary axes of trait covariation. The first exhibited differences between species with dense tissues and species with soft tissues (the tissue construction cost axis); however, the oak sections did not constitute separate groups, while the second suggested a trade-off between xylem resistance to cavitation and tree deciduousness. As expected, the water-use strategies of the species were related to the environment; oak species from arid areas had more deciduousness and a higher instantaneous water-use efficiency. In contrast, their humid counterparts had less deciduousness and had a xylem that was more resistant to embolisms. Altogether, these results suggest that aridity filters closely related species, resulting in habitat partitioning and niche divergence.

Exploiting water versus tolerating drought: water-use strategies of trees in a secondary successional tropical dry forest.

In seasonal plant communities where water availability changes dramatically both between and within seasons, understanding the mechanisms that enable plants to exploit water pulses and to survive drought periods is crucial. By measuring rates of physiological processes, we examined the trade-off between water exploitation and drought tolerance among seedlings of trees of a tropical dry forest, and identified biophysical traits most closely associated with plant water-use strategies. We also explored whether early and late secondary successional species occupy different portions of trade-off axes. As predicted, species that maintained carbon capture, hydraulic function and leaf area at higher plant water deficits during drought had low photosynthetic rates, xylem hydraulic conductivity and growth rate under non-limiting water supply. Drought tolerance was associated with more dense leaf, stem and root tissues, whereas rapid resource acquisition was associated with greater stem water storage, larger vessel diameter and larger leaf area per mass invested. We offer evidence that the water exploitation versus drought tolerance trade-off drives species differentiation in the ability of tropical dry forest trees to deal with alternating water-drought pulses. However, we detected no evidence of strong functional differentiation between early and late successional species along the proposed trade-off axes, suggesting that the environmental gradient of water availability across secondary successional habitats in the dry tropics does not filter out physiological strategies of water use among species, at least at the seedling stage.

Root depth and morphology in response to soil drought: comparing ecological groups along the secondary succession in a tropical dry forest.

Root growth and morphology may play a core role in species-niche partitioning in highly diverse communities, especially along gradients of drought risk, such as that created along the secondary succession of tropical dry forests. We experimentally tested whether root foraging capacity, especially at depth, decreases from early successional species to old-growth forest species. We also tested for a trade-off between two mechanisms for delaying desiccation, the capacity to forage deeper in the soil and the capacity to store water in tissues, and explored whether successional groups separate along such a trade-off. We examined the growth and morphology of roots in response to a controlled-vertical gradient of soil water, among seedlings of 23 woody species dominant along the secondary succession in a tropical dry forest of Mexico. As predicted, successional species developed deeper and longer root systems than old-growth forest species in response to soil drought. In addition, shallow root systems were associated with high plant water storage and high water content per unit of tissue in stems and roots, while deep roots exhibited the opposite traits, suggesting a trade-off between the capacities for vertical foraging and water storage. Our results suggest that an increased capacity of roots to forage deeper for water is a trait that enables successional species to establish under the warm-dry conditions of the secondary succession, while shallow roots, associated with a higher water storage capacity, are restricted to the old-growth forest. Overall, we found evidence that the root depth-water storage trade-off may constrain tree species distribution along secondary succession.

Drought resistance in early and late secondary successional species from a tropical dry forest: the interplay between xylem resistance to embolism, sapwood water storage and leaf shedding.

The mechanisms of drought resistance that allow plants to successfully establish at different stages of secondary succession in tropical dry forests are not well understood. We characterized mechanisms of drought resistance in early and late-successional species and tested whether risk of drought differs across sites at different successional stages, and whether early and late-successional species differ in resistance to experimentally imposed soil drought. The microenvironment in early successional sites was warmer and drier than in mature forest. Nevertheless, successional groups did not differ in resistance to soil drought. Late-successional species resisted drought through two independent mechanisms: high resistance of xylem to embolism, or reliance on high stem water storage capacity. High sapwood water reserves delayed the effects of soil drying by transiently decoupling plant and soil water status. Resistance to soil drought resulted from the interplay between variations in xylem vulnerability to embolism, reliance on sapwood water reserves and leaf area reduction, leading to a tradeoff of avoidance against tolerance of soil drought, along which successional groups were not differentiated. Overall, our data suggest that ranking species' performance under soil drought based solely on xylem resistance to embolism may be misleading, especially for species with high sapwood water storage capacity.

Morphological and physiological differentiation of seedlings between dry and wet habitats in a tropical dry forest.

A common observation in tropical dry forests is the habitat preference of tree species along spatial soil water gradients. This pattern of habitat partitioning might be a result of species differentiation in their strategy for using water, along with competing functions such as maximizing water exploitation and tolerating soil water stress. We tested whether species from drier soil conditions exhibited a tolerance strategy compared with that of wet-habitat species. In a comparison of 12 morphophysiological traits in seedlings of 10 closely related dry and wet-habitat species pairs, we explored what trade-offs guide differentiation between habitats and species. Contrary to our expectations, dry-habitat species showed mostly traits associated with an exploitation strategy (higher carbon assimilation capacity, specific leaf area and leaf-specific conductivity and lower water-use efficiency). Strikingly, dry-habitat species tended to retain their leaves longer during drought. Additionally, we detected multiple strategies to live within each habitat, in part due to variation of strategies among lineages, as well as functional differentiation along the water storage capacity-stem density (xylem safety) trade-off. Our results suggest that fundamental trade-offs guide functional niche differentiation among tree species expressed both within and between soil water habitats in a tropical dry forest.