Summary The temperature response of photosynthesis is one of the key factors determining predicted responses to warming in global vegetation models (GVMs). The response may vary geographically, owing to genetic adaptation to climate, and temporally, as a result of acclimation to changes in ambient temperature. Our goal was to develop a robust quantitative global model representing acclimation and adaptation of photosynthetic temperature responses. We quantified and modelled key mechanisms responsible for photosynthetic temperature acclimation and adaptation using a global dataset of photosynthetic CO2 response curves, including data from 141 C3 species from tropical rainforest to Arctic tundra. We separated temperature acclimation and adaptation processes by considering seasonal and common‐garden datasets, respectively. The observed global variation in the temperature optimum of photosynthesis was primarily explained by biochemical limitations to photosynthesis, rather than stomatal conductance or respiration. We found acclimation to growth temperature to be a stronger driver of this variation than adaptation to temperature at climate of origin. We developed a summary model to represent photosynthetic temperature responses and showed that it predicted the observed global variation in optimal temperatures with high accuracy. This novel algorithm should enable improved prediction of the function of global ecosystems in a warming climate.

Although tropical forests account for only a fraction of the planet's 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.

ABSTRACT[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.

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.

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 (CO(2) ) 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.

The balance between photosynthesis and plant respiration in tropical forests may substantially affect the global carbon cycle. Woody tissue CO 2 efflux is a major component of total plant respiration, but estimates of ecosystem-scale rates are uncertain because of poor sampling in the upper canopy and across landscapes. To overcome these problems, we used a portable scaffolding tower to measure woody tissue CO 2 efflux from ground level to the canopy top across a range of sites of varying slope and soil phosphorus content in a primary tropical rain forest in Costa Rica. The objectives of this study were to: (1) determine whether to use surface area, volume, or biomass for modeling and extrapolating wood CO 2 efflux, (2) determine if wood CO 2 efflux varied seasonally, (3) identify if wood CO 2 efflux varied by functional group, height in canopy, soil fertility, or slope, and (4) extrapolate wood CO 2 efflux to the forest. CO 2 efflux from small diameter woody tissue (o10 cm) was related to surface area, while CO 2 efflux from stems 410 cm was related to both surface area and volume. Wood CO 2 efflux showed no evidence of seasonality over 2 years. CO 2 efflux per unit wood surface area at 251 (F A ) was highest for the N-fixing dominant tree species Pentaclethra macroloba, followed by other tree species, lianas, then palms. Small diameter F A increased steeply with increasing height, and large diameter F A increased with diameter. Soil phosphorus and slope had slight, but complex effects on F A . Wood CO 2 efflux per unit ground area was 1.34 AE 0.36 lmol m À2 s À1 , or 508 AE 135 g C m À2 yr À1 . Small diameter wood, only 15% of total woody biomass, accounted for 70% of total woody tissue CO 2 efflux from the forest; while lianas, only 3% of total woody biomass, contributed one-fourth of the total wood CO 2 efflux.

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.

The response of tropical forests to global warming is one of the largest uncertainties in predicting the future carbon balance of Earth. To determine the likely effects of elevated temperatures on tropical forest understory plants and soils, as well as other ecosystems, an infrared (IR) heater system was developed to provide in situ warming for the Tropical Responses to Altered Climate Experiment (TRACE) in the Luquillo Experimental Forest in Puerto Rico. Three replicate heated 4‐m‐diameter plots were warmed to maintain a 4°C increase in understory vegetation compared to three unheated control plots, as sensed by IR thermometers. The equipment was larger than any used previously and was subjected to challenges different from those of many temperate ecosystem warming systems, including frequent power surges and outages, high humidity, heavy rains, hurricanes, saturated clayey soils, and steep slopes. The system was able to maintain the target 4.0°C increase in hourly average vegetation temperatures to within ± 0.1°C. The vegetation was heterogeneous and on a 21° slope, which decreased uniformity of the warming treatment on the plots; yet, the green leaves were fairly uniformly warmed, and there was little difference among 0–10 cm depth soil temperatures at the plot centers, edges, and midway between. Soil temperatures at the 40–50 cm depth increased about 3°C compared to the controls after a month of warming. As expected, the soil in the heated plots dried faster than that of the control plots, but the average soil moisture remained adequate for the plants. The TRACE heating system produced an adequately uniform warming precisely controlled down to at least 50‐cm soil depth, thereby creating a treatment that allows for assessing mechanistic responses of tropical plants and soil to warming, with applicability to other ecosystems. No physical obstacles to scaling the approach to taller vegetation (i.e., trees) and larger plots were observed.