Molly A. Cavaleri

Comparative water use of native and invasive plants at multiple scales: a global meta‐analysis

Ecohydrology and invasive ecology have become increasingly important in the context of global climate change. This study presents the first in-depth analysis of the water use of invasive and native plants of the same growth form at multiple scales: leaf, plant, and ecosystem. We reanalyzed data for several hundred native and invasive species from over 40 published studies worldwide to glean global trends and to highlight how patterns vary depending on both scale and climate. We analyzed all pairwise combinations of co-occurring native and invasive species for higher comparative resolution of the likelihood of an invasive species using more water than a native species and tested for significance using bootstrap methods. At each scale, we found several-fold differences in water use between specific paired invasive and native species. At the leaf scale, we found a strong tendency for invasive species to have greater stomatal conductance than native species. At the plant scale, however, natives and invasives were equally likely to have the higher sap flow rates. Available data were much fewer for the ecosystem scale; nevertheless, we found that invasive-dominated ecosystems were more likely to have higher sap flow rates per unit ground area than native-dominated ecosystems. Ecosystem-scale evapotranspiration, on the other hand, was equally likely to be greater for systems dominated by invasive and native species of the same growth form. The inherent disconnects in the determination of water use when changing scales from leaf to plant to ecosystem reveal hypotheses for future studies and a critical need for more ecosystem-scale water use measurements in invasive- vs. native-dominated systems. The differences in water use of native and invasive species also depended strongly on climate, with the greater water use of invasives enhanced in hotter, wetter climates at the coarser scales.

Height is more important than light in determining leaf morphology in a tropical forest

Both within and between species, leaf physiological parameters are strongly related to leaf dry mass per area (LMA, g/m2), which has been found to increase from forest floor to canopy top in every forest where it has been measured. Although vertical LMA gradients in forests have historically been attributed to a direct phenotypic response to light, an increasing number of recent studies have provided evidence that water limitation in the upper canopy can constrain foliar morphological adaptations to higher light levels. We measured height, light, and LMA of all species encountered along 45 vertical canopy transects across a Costa Rican tropical rain forest. LMA was correlated with light levels in the lower canopy until approximately 18 m sample height and 22% diffuse transmittance. Height showed a remarkably linear relationship with LMA throughout the entire vertical canopy profile for all species pooled and for each functional group individually (except epiphytes), possibly through the influence of gravity on leaf water potential and turgor pressure. Models of forest function may be greatly simplified by estimating LMA-correlated leaf physiological parameters solely from foliage height profiles, which in turn can be assessed with satellite- and aircraft-based remote sensing.

Foliar and ecosystem respiration in an old-growth tropical rain forest

Foliar respiration is a major component of ecosystem respiration, yet extrapolations are often uncertain in tropical forests because of indirect estimates of leaf area index (LAI). A portable tower was used to directly measure LAI and night-time foliar respiration from 52 vertical transects throughout an old-growth tropical rain forest in Costa Rica. In this study, we (1) explored the effects of structural, functional and environmental variables on foliar respiration; (2) extrapolated foliar respiration to the ecosystem; and (3) estimated ecosystem respiration. Foliar respiration temperature response was constant within plant functional group, and foliar morphology drove much of the within-canopy variability in respiration and foliar nutrients. Foliar respiration per unit ground area was 3.5 +/- 0.2 micromol CO2 m(-2) s(-1), and ecosystem respiration was 9.4 +/- 0.5 micromol CO2 m(-2) s(-1)[soil = 41%; foliage = 37%; woody = 14%; coarse woody debris (CWD) = 7%]. When modelled with El Niño Southern Oscillation (ENSO) year temperatures, foliar respiration was 9% greater than when modelled with temperatures from a normal year, which is in the range of carbon sink versus source behaviour for this forest. Our ecosystem respiration estimate from component fluxes was 33% greater than night-time net ecosystem exchange for the same forest, suggesting that studies reporting a large carbon sink for tropical rain forests based solely on eddy flux measurements may be in error.

Tropical forest carbon balance in a warmer world: a critical review spanning microbial‐ to ecosystem‐scale processes

Tropical forests play a major role in regulating global carbon (C) fluxes and stocks, and even small changes to C cycling in this productive biome could dramatically affect atmospheric carbon dioxide (CO2) concentrations. Temperature is expected to increase over all land surfaces in the future, yet we have a surprisingly poor understanding of how tropical forests will respond to this significant climatic change. Here we present a contemporary synthesis of the existing data and what they suggest about how tropical forests will respond to increasing temperatures. Our goals were to: (i) determine whether there is enough evidence to support the conclusion that increased temperature will affect tropical forest C balance; (ii) if there is sufficient evidence, determine what direction this effect will take; and, (iii) establish what steps should to be taken to resolve the uncertainties surrounding tropical forest responses to increasing temperatures. We approach these questions from a mass‐balance perspective and therefore focus primarily on the effects of temperature on inputs and outputs of C, spanning microbial‐ to ecosystem‐scale responses. We found that, while there is the strong potential for temperature to affect processes related to C cycling and storage in tropical forests, a notable lack of data combined with the physical, biological and chemical diversity of the forests themselves make it difficult to resolve this issue with certainty. We suggest a variety of experimental approaches that could help elucidate how tropical forests will respond to warming, including large‐scale in situ manipulation experiments, longer term field experiments, the incorporation of a range of scales in the investigation of warming effects (both spatial and temporal), as well as the inclusion of a diversity of tropical forest sites. Finally, we highlight areas of tropical forest research where notably few data are available, including temperature effects on: nutrient cycling, heterotrophic versus autotrophic respiration, thermal acclimation versus substrate limitation of plant and microbial communities, below‐ground C allocation, species composition (plant and microbial), and the hydraulic architecture of roots. Whether or not tropical forests will become a source or a sink of C in a warmer world remains highly uncertain. Given the importance of these ecosystems to the global C budget, resolving this uncertainty is a primary research priority.

Urgent need for warming experiments in tropical forests

Although tropical forests account for only a fraction of the planets terrestrial surface, they exchange more carbon dioxide with the atmosphere than any other biome on Earth, and thus play a disproportionate role in the global climate. In the next 20 years, the tropics will experience unprecedented warming, yet there is exceedingly high uncertainty about their potential responses to this imminent climatic change. Here, we prioritize research approaches given both funding and logistical constraints in order to resolve major uncertainties about how tropical forests function and also to improve predictive capacity of earth system models. We investigate overall model uncertainty of tropical latitudes and explore the scientific benefits and inevitable trade-offs inherent in large-scale manipulative field experiments. With a Coupled Model Intercomparison Project Phase 5 analysis, we found that model variability in projected net ecosystem production was nearly 3 times greater in the tropics than for any other latitude. Through a review of the most current literature, we concluded that manipulative warming experiments are vital to accurately predict future tropical forest carbon balance, and we further recommend the establishment of a network of comparable studies spanning gradients of precipitation, edaphic qualities, plant types, and/or land use change. We provide arguments for long-term, single-factor warming experiments that incorporate warming of the most biogeochemically active ecosystem components (i.e. leaves, roots, soil microbes). Hypothesis testing of underlying mechanisms should be a priority, along with improving model parameterization and constraints. No single tropical forest is representative of all tropical forests; therefore logistical feasibility should be the most important consideration for locating large-scale manipulative experiments. Above all, we advocate for multi-faceted research programs, and we offer arguments for what we consider the most powerful and urgent way forward in order to improve our understanding of tropical forest responses to climate change.

Wood CO2 efflux and foliar respiration for Eucalyptus in Hawaii and Brazil

We measured CO(2) efflux from wood for Eucalyptus in Hawaii for 7 years and compared these measurements with those on three- and four-and-a-half-year-old Eucalyptus in Brazil. In Hawaii, CO(2) efflux from wood per unit biomass declined approximately 10x from age two to age five, twice as much as the decline in tree growth. The CO(2) efflux from wood in Brazil was 8-10x lower than that for comparable Hawaii trees with similar growth rates. Growth and maintenance respiration coefficients calculated from Hawaii wood CO(2) efflux declined with tree age and size (the growth coefficient declined from 0.4 mol C efflux mol C(-1) wood growth at age one to 0.1 mol C efflux mol C(-1) wood growth at age six; the maintenance coefficient from 0.006 to 0.001 micromol C (mol C biomass)(-1) s(-1) at 20 degrees C over the same time period). These results suggest interference with CO(2) efflux through bark that decouples CO(2) efflux from respiration. We also compared the biomass fractions and wood CO(2) efflux for the aboveground woody parts for 3- and 7-year-old trees in Hawaii to estimate how focusing measurements near the ground might bias the stand-level estimates of wood CO(2) efflux. Three-year-old Eucalyptus in Hawaii had a higher proportion of branches < 0.5 cm in diameter and a lower proportion of stem biomass than did 7-year-old trees. Biomass-specific CO(2) efflux measured at 1.4 m extrapolated to the tree could bias tree level estimates by approximately 50%, assuming no refixation from bark photosynthesis. However, the bias did not differ for the two tree sizes. Foliar respiration was identical per unit nitrogen for comparable treatments in Brazil and Hawaii (4.2 micromol C mol N(-1) s(-1) at 20 degrees C).

Light acclimation optimizes leaf functional traits despite height-related constraints in a canopy shading experiment.

Within-canopy gradients of leaf functional traits have been linked to both light availability and vertical gradients in leaf water potential. While observational studies can reveal patterns in leaf traits, within-canopy experimental manipulations can provide mechanistic insight to tease apart multiple interacting drivers. Our objectives were to disentangle effects of height and light environment on leaf functional traits by experimentally shading branches along vertical gradients within a sugar maple (Acer saccharum) forest. Shading reduced leaf mass per area (LMA), leaf density, area-based leaf nitrogen (Narea), and carbon:nitrogen (C:N) ratio, and increased mass-based leaf nitrogen (Nmass), highlighting the importance of light availability on leaf morphology and chemistry. Early in the growing season, midday leaf water potential (Ψmid), LMA, and Narea were driven primarily by height; later in the growing season, light became the most important driver for LMA and Narea. Carbon isotope composition (δ13C) displayed strong, linear correlations with height throughout the growing season, but did not change with shading, implying that height is more influential than light on water use efficiency and stomatal behavior. LMA, leaf density, Nmass, C:N ratio, and δ13C all changed seasonally, suggesting that leaf ageing effects on leaf functional traits are equally as important as microclimatic conditions. Overall, our results indicate that: (1) stomatal sensitivity to vapor pressure deficit or Ψmid constrains the supply of CO2 to leaves at higher heights, independent of light environment, and (2) LMA and Narea distributions become functionally optimized through morphological acclimation to light with increasing leaf age despite height-related constraints.

Light drives vertical gradients of leaf morphology in a sugar maple (Acer saccharum) forest

Leaf mass per area (LMA, g m(-2)) is an essential trait for modeling canopy function due to its strong association with photosynthesis, respiration and leaf nitrogen. Leaf mass per area, which is influenced by both leaf thickness and density (LMA = thickness × density), generally increases from the bottom to the top of tree canopies, yet the mechanisms behind this universal pattern are not yet resolved. For decades, the light environment was assumed to be the most influential driver of within-canopy variation in LMA, yet recent evidence has shown hydrostatic gradients to be more important in upper canopy positions, especially in tall evergreen trees in temperate and tropical forests. The aim of this study was to disentangle the importance of various environmental drivers on vertical LMA gradients in a mature sugar maple (Acer saccharum Marshall) forest. We compared LMA, leaf density and leaf thickness relationships with height, light and predawn leaf water potential (ΨPre) within a closed and an exposed canopy to assess leaf morphological traits at similar heights but different light conditions. Contrary to our expectations and recent findings in the literature, we found strong evidence that light was the primary driver of vertical gradients in leaf morphology. At similar heights (13-23 m), LMA was greater within the exposed canopy than the closed canopy, and light had a stronger influence over LMA compared with ΨPre. Light also had a stronger influence over both leaf thickness and density compared with ΨPre; however, the increase in LMA within both canopy types was primarily due to increasing leaf thickness with increasing light availability. This study provides strong evidence that canopy structure and crown exposure, in addition to height, should be considered as a parameter for determining vertical patterns in LMA and modeling canopy function.

Native trees show conservative water use relative to invasive trees: results from a removal experiment in a Hawaiian wet forest

To gain understanding about invasive species physiology and also potential water conservation strategies, we investigated tree-level water use following a woody invasive removal experiment in Hawaii. Invasives had much higher water use rates than the native tree species, and stand level water use decreased by ∼50% after invasive removal.

Tropical forests in a warming world.

Global models suggest tropical forests could face significant and unprecedented warming within the next two decades (Diffenbaugh & Scherer, 2011). These findings, combined with evidence that tropical forests may be near a high temperature threshold, suggest that these systems may be more vulnerable to climate change than previously believed (Clark et al., 2003; Doughty & Goulden, 2008). Currently, our ability to predict tropical forest responses to rising temperatures is limited, due in large part to a lack of data on tropical forest and tree processes (Saxe et al., 2001). Given the immense amount of carbon (C) cycled through these forests, even a slight change to tropical net C fluxes could have significant consequences for global C cycling and future climate. The organized oral session From Leaf to Biosphere: The Effects of a Warming Climate on Tropical Forests at the 2011 Ecological Society of America meeting in Austin, Texas brought together 10 scientists from a range of fields to synthesize existing data on tropical forest responses to increasing temperature and to facilitate cross-disciplinary dialogue. These scientists presented research and offered perspectives spanning molecular to global scales (Fig. 1). Here, we synthesize the key conclusions.