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  1. Free, publicly-accessible full text available June 1, 2024
  2. Abstract

    Root‐based functional traits are relatively overlooked as drivers of savanna plant community dynamics, an important gap in water‐limited ecosystems. Recent work has shed light on patterns of trait coordination in roots, but less is known about the relationship between root functional traits, water acquisition, and plant demographic rates. Here, we investigated how fine‐root vascular and morphological traits are related in two dominant PFTs (C3trees and C4grasses from the savanna biome), whether root traits can predict plant relative growth rate (RGR), and whether root trait multivariate relationships differ in trees and grasses. We used root data from 21 tree and 18 grass species grown under greenhouse conditions, and quantified a suite of vascular and morphological root traits. We used a principal components analysis (PCA) to identify common axes of trait variation, compared trait correlation matrices between the two PFTs, and investigated the relationship between PCA axes and individual traits and RGR. We found that there was no clear single axis integrating vascular and morphological traits, but found that vascular anatomy predicted RGR in both trees and grasses. Trait correlation matrices differed in trees and grasses, suggesting potentially divergent patterns of trait coordination between the two functional types. Our results suggested that, despite differences in trait relationships between trees and grasses, root conductivity may constrain maximum growth rate in both PFTs, highlighting the critical role that water relations play in savanna vegetation dynamics and suggesting that root water transport capacity is an important predictor of plant performance in the savanna biome.

     
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  3. Summary

    Savannas cover a significant fraction of the Earth's land surface. In these ecosystems, C3trees and C4grasses coexist persistently, but the mechanisms explaining coexistence remain subject to debate. Different quantitative models have been proposed to explain coexistence, but these models make widely contrasting assumptions about which mechanisms are responsible for savanna persistence. Here, we show that no single existing model fully captures all key elements required to explain tree–grass coexistence across savanna rainfall gradients, but many models make important contributions. We show that recent empirical work allows us to combine many existing elements with new ideas to arrive at a synthesis that combines elements of two dominant frameworks: Walter's two‐layer model and demographic bottlenecks. We propose that functional rooting separation is necessary for coexistence and is the crux of the coexistence problem. It is both well‐supported empirically and necessary for tree persistence, given the comprehensive grass superiority for soil moisture acquisition. We argue that eventual tree dominance through shading is precluded by ecohydrological constraints in dry savannas and by fire and herbivores in wet savannas. Strong asymmetric grass–tree competition for soil moisture limits tree growth, exposing trees to persistent demographic bottlenecks.

     
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  4. Summary

    Models of tree–grass coexistence in savannas make different assumptions about the relative performance of trees and grasses under wet vs dry conditions. We quantified transpiration and drought tolerance traits in 26 tree and 19 grass species from the African savanna biome across a gradient of soil water potentials to test for a trade‐off between water use under wet conditions and drought tolerance.

    We measured whole‐plant hourly transpiration in a growth chamber and quantified drought tolerance using leaf osmotic potential (Ψosm). We also quantified whole‐plant water‐use efficiency (WUE) and relative growth rate (RGR) under well‐watered conditions.

    Grasses transpired twice as much as trees on a leaf‐mass basis across all soil water potentials. Grasses also had a lower Ψosmthan trees, indicating higher drought tolerance in the former. Higher grass transpiration and WUE combined to largely explain the threefold RGR advantage in grasses.

    Our results suggest that grasses outperform trees under a wide range of conditions, and that there is no evidence for a trade‐off in water‐use patterns in wet vs dry soils. This work will help inform mechanistic models of water use in savanna ecosystems, providing much‐needed whole‐plant parameter estimates for African species.

     
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  5. Premise

    Belowground functional traits play a significant role in determining plant water‐use strategies and plant performance, but we lack data on root traits across communities, particularly in the tropical savanna biome, where vegetation dynamics are hypothesized to be strongly driven by tree–grass functional differences in water use.

    Methods

    We grew seedlings of 21 tree and 18 grass species (N= 5 individuals per species) from the southern African savanna biome under greenhouse conditions and collected fine‐root segments from plants for histological analysis. We identified and measured xylem vessels in 539 individual root cross sections. We then quantified six root vascular anatomy traits and tested them for phylogenetic signals and tree–grass differences in trait values associated with vessel size, number, and hydraulic conductivity.

    Results

    Grass roots had larger root xylem vessels than trees, a higher proportion of their root cross‐sectional area comprised vessels, and they had higher estimated axial conductivities than trees, while trees had a higher number of vessels per root cross‐sectional area than grasses did. We found evidence of associations between trait values and phylogenetic relatedness in most of these traits across tree species, but not grasses.

    Conclusions

    Our findings support the hypothesis that grass roots have higher water transport capacity than tree roots in terms of maximum axial conductivity, consistent with the observation that grasses are more “aggressive” water users than trees under conditions of high soil moisture availability. Our study identifies root functional traits that may drive differential responses of trees and grasses to soil moisture availability.

     
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  6. Abstract

    In savannas, partitioning of below‐ground resources by depth could facilitate tree–grass coexistence and shape vegetation responses to changing rainfall patterns. However, most studies assessing tree versus grass root‐niche partitioning have focused on one or two sites, limiting generalization about how rainfall and soil conditions influence the degree of rooting overlap across environmental gradients.

    We used two complementary stable isotope techniques to quantify variation (a) in water uptake depths and (b) in fine‐root biomass distributions among dominant trees and grasses at eight semi‐arid savanna sites in Kruger National Park, South Africa. Sites were located on contrasting soil textures (clayey basaltic soils vs. sandy granitic soils) and paired along a gradient of mean annual rainfall.

    Soil texture predicted variation in mean water uptake depths and fine‐root allocation. While grasses maintained roots close to the surface and consistently used shallow water, trees on sandy soils distributed roots more evenly across soil depths and used deeper soil water, resulting in greater divergence between tree and grass rooting on sandy soils. Mean annual rainfall predicted some variation among sites in tree water uptake depth, but had a weaker influence on fine‐root allocation.

    Synthesis. Savanna trees overlapped more with shallow‐rooted grasses on clayey soils and were more distinct in their use of deeper soil layers on sandy soils, consistent with expected differences in infiltration and percolation. These differences, which could allow trees to escape grass competition more effectively on sandy soils, may explain observed differences in tree densities and rates of woody encroachment with soil texture. Differences in the degree of root‐niche separation could also drive heterogeneous responses of savanna vegetation to predicted shifts in the frequency and intensity of rainfall.

     
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  7. Abstract

    Although there is a well‐known association between tree cover and soil texture in savannahs, the hydrological drivers of tree cover variation have not been systematically explored, particularly in parallel with factors such as fire, herbivory, and tree–grass interactions. The relationship between hydrological factors and tree cover is important for resolving the relative contribution of bottom‐up versus top‐down factors in structuring savannah vegetation.

    We quantified soil moisture dynamics across eight 1‐km transects spanning tree cover gradients from open to woody savannah in Serengeti National Park in Tanzania using soil moisture sensors coupled with dataloggers. We mapped tree cover at two spatial scales through supervised classification of high‐resolution satellite imagery. We simultaneously produced water retention curves in open and woody habitats within each transect to compare soil hydrological properties and to convert volumetric water content (θ) from dataloggers to plant‐available water over the course of an annual cycle. We also quantified grass biomass at 100 locations per transect, estimated fire frequency from MODIS satellite data, and quantified herbivore occupancy with paired camera traps situated in open and woody habitats within each transect.

    We found a positive relationship between tree cover and soil moisture drainage rate, and found that open habitats had more negative water potentials than woody habitats for a given value ofθ. In contrast, we found no evidence for a consistent relationship between grass biomass or fire frequency and tree cover. We found evidence for higher browser occupancy in woody than open habitats, but no habitat effects on herbivores as a group (browsers plus grazers), suggesting that herbivory is unlikely to be the dominant factor explaining variation in tree cover.

    Synthesis. Our results suggest that variation in tree cover is partly driven by hydrological (edaphic) factors unrelated to fire, herbivory, tree–grass interactions or mean annual precipitation at these spatial scales in Serengeti. We contrast our findings with previous work attributing tree cover shifts in Serengeti to precipitation gradients.

     
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  8. Abstract

    Savanna tree species vary in the magnitude of their response to grass competition, but the functional traits that explain this variation remain largely unknown. To address this gap, we grew seedlings of 10 savanna tree species with and without grasses in a controlled greenhouse experiment. We found strong interspecific differences in tree competitive response, which was positively related to photosynthesis rates, suggesting a trade‐off between the ability to grow well under conditions of low and high grass biomass across tree species. We also found no competitive effect of tree seedlings on grass, suggesting strong tree‐grass competitive asymmetry. Our results identify a potentially important trade‐off that enhances our ability to predict how savanna tree communities might respond to variation in grass competition.

     
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