• Tropical climates are getting warmer, with pronounced dry periods in large areas. The productivity and climate feedbacks of future tropical forests depend on the ability of trees to acclimate their physiological processes, such as leaf dark respiration (R d), to these new conditions. However, knowledge on this is currently limited due to data scarcity. • We studied the impact of growth temperature on R d and its dependency on net photosynthesis (A n), leaf nitrogen (N) and phosphorus (P) contents, and leaf mass per unit area (LMA) in 16 early-(ES) and late-successional (LS) tropical tree species in multi-species plantations along an elevation gradient. Moreover, we explored the effect of drought on R d in one ES and one LS species. • Leaf R d at 20 °C decreased at warmer sites, regardless if it was expressed per unit leaf area, mass, N or P. This acclimation resulted in 8% and 28% decrease in R d at prevailing nighttime temperatures in trees at the intermediate and warmest sites, respectively. Moreover, drought reduced R d , particularly in the ES species and at the coolest site. • Thermal acclimation of R d is complete or over-compensatory and independent of changes in leaf nutrients or LMA in African tropical trees.
Warming climate increases the risk for harmful leaf temperatures in terrestrial plants, causing heat stress and loss of productivity. The heat sensitivity may be particularly high in equatorial tropical tree species adapted to a thermally stable climate.Thermal thresholds of the photosynthetic system of sun-exposed leaves were investigated in three tropical montane tree species native to Rwanda with different growth and water use strategies (Harungana montana, Syzygium guineense and Entandrophragma exselsum). Measurements of chlorophyll fluorescence, leaf gas exchange, morphology, chemistry and temperature were made at three common gardens along an elevation/temperature gradient.Heat tolerance acclimated to maximum leaf temperature (T leaf ) across the species. At the warmest sites, the thermal threshold for normal function of photosystem II was exceeded in the species with the highest T leaf despite their higher heat tolerance. This was not the case in the species with the highest transpiration rates and lowest T leaf . The results point to two differently effective strategies for managing thermal stress: tolerance through physiological adjustment of leaf osmolality and thylakoid membrane lipid composition, or avoidance through morphological adaptation and transpiratory cooling.More severe photosynthetic heat stress in low-transpiring montane climax species may result in a competitive disadvantage compared to high-transpiring pioneer species with more efficient leaf cooling.
Leaf morphological traits vary along climate gradients, but it is currently unclear to what extent this results from acclimation rather than adaptation. Knowing so is important for predicting the functioning of long-lived organisms, such as trees, in a rapidly changing climate. We investigated the leaf morphological warming responses of 18 tropical tree species with early (ES) abd late (LS) successional strategies, planted at three sites along an elevation gradient from 2400 m a.s.l. (15.2 °C mean temperature) to 1300 m a.s.l. (20.6 °C mean temperature) in Rwanda. Leaf size expressed as leaf area (LA) and leaf mass per area (LMA) decreased, while leaf width-to-length ratio (W/L) increased with warming, but only for one third to half of the species. While LA decreased in ES species, but mostly not in LS species, changes in LMA and leaf W/L were common in both successional groups. ES species had lower LMA and higher LA and leaf W/L compared to LS species. Values of LMA and LA of juvenile trees in this study were mostly similar to corresponding data on four mature tree species in another elevation-gradient study in Rwanda, indicating that our results are applicable also to mature forest trees. We conclude that leaf morphological responses to warming differ greatly between both successional groups and individual species, with potential consequences for species competitiveness and community composition in a warmer climate.
Elevation gradients offer excellent opportunities to explore the climate sensitivity of vegetation. Here, we investigated elevation patterns of structural, chemical, and physiological traits in tropical tree species along a 1700–2700 m elevation gradient in Rwanda, central Africa. Two early-successional (Polyscias fulva, Macaranga kilimandscharica) and two late-successional (Syzygium guineense, Carapa grandiflora) species that are abundant in the area and present along the entire gradient were investigated. We found that elevation patterns in leaf stomatal conductance (gs), transpiration (E), net photosynthesis (An), and water-use efficiency were highly season-dependent. In the wet season, there was no clear variation in gs or An with elevation, while E was lower at cooler high-elevation sites. In the dry season, gs, An, and E were all lower at drier low elevation sites. The leaf-to-air temperature difference was smallest in P. fulva, which also had the highest gs and E. Water-use efficiency (An/E) increased with elevation in the wet season, but not in the dry season. Leaf nutrient ratios indicated that trees at all sites are mostly P limited and the N:P ratio did not decrease with increasing elevation. Our finding of strongly decreased gas exchange at lower sites in the dry season suggests that both transpiration and primary production would decline in a climate with more pronounced dry periods. Furthermore, we showed that N limitation does not increase with elevation in the forests studied, as otherwise most commonly reported for tropical montane forests.
Abstract. The response of tropical trees and tree communities to climate change is crucial for the carbon storage and biodiversity of the terrestrial biosphere. Trees in tropical montane rainforests (TMFs) are considered particularly vulnerable to climate change, but this hypothesis remains poorly evaluated due to data scarcity. To reduce the knowledge gap on the response of TMFs trees to warming, we established a field experiment along a 1300–2400 m elevation gradient in Rwanda. Twenty tree species native to montane forests in East and Central Africa were planted in multispecies plots at three sites along the gradient. They have overlapping distributions but primarily occur in either transitional rainforest (1600–2000 m a.s.l) or mid elevation TMF (2000–3000 m a.s.l.), with both early- (ES) and late-successional (LS) species represented in each elevation origin group. Tree growth (diameter and height) and survival were monitored regularly over two years. We found that ES species, especially from lower elevations, grew faster at warmer sites while several of the LS species, especially from higher elevations, did not respond or grew slower. Moreover, a warmer climate increased tree mortality in LS species, but not much in ES species. ES species with transitional rainforest origin strongly increased in proportion of stand basal area at warmer sites, while TMF species declined, suggesting that lower-elevation ES species will have an advantage over higher-elevation species in a warming climate. The risk of higher-elevation and LS species to become outcompeted by lower-elevation and ES species in a warmer climate has important implications for biodiversity and carbon storage of Afromontane forests.
<p>Current estimates of temperature effects on plants are usually based on air temperature (<em>T</em><sub>air</sub>), although it is well known that leaf temperature (<em>T</em><sub>leaf</sub>) can deviate considerably from <em>T</em><sub>air</sub>. In some studies, to overcome the problem of <em>T</em><sub>air</sub> often being a poor proxy of <em>T</em><sub>leaf</sub>, measurements of canopy temperature (<em>T</em><sub>can</sub>) have been used instead. However, <em>T</em><sub>can</sub> data do not capture the spatial variation in <em>T</em><sub>leaf</sub> among leaves with different thermoregulatory traits. This may be particularly problematic for highly diverse and heterogeneous tropical forest canopies. In this study, we used infrared thermometers to study <em>T</em><sub>leaf</sub> and <em>T</em><sub>can</sub> in multispecies tropical tree plantations established at three sites along an elevation gradient from 2,400 m a.s.l. (17.1&#176;C mean daytime temperature) to 1,300 m a.s.l. (24.0&#176;C) in Rwanda. &#160;Measurements of chlorophyll fluorescence were also conducted to study the photosynthetic heat tolerance of these species. Our results showed high <em>T</em><sub>leaf</sub> (up to ~50&#176;C) and leaf-to-air temperature differences (&#916;<em>T</em><sub>leaf</sub>; on average 8-10&#176;C and up to 24&#176;C) of sun-exposed leaves. Both leaf size and stomatal conductance were important traits in controlling <em>T</em><sub>leaf</sub>. The <em>T</em><sub>leaf</sub> (and thus &#916;<em>T</em><sub>leaf</sub>) of sun-exposed leaves greatly exceeded the simultaneously measured values of <em>T</em><sub>can</sub> (and &#916;<em>T</em><sub>can</sub>). Photosynthetic heat tolerance partially acclimated to increased growth temperature; on average 0.31&#176;C increase in heat tolerance per 1&#176;C increase in growth temperature. Consequently, thermal safety margins were narrower for species at the warmer, lower-elevation sites. Our findings highlight the importance of leaf traits for leaf thermoregulation and show that monitoring of canopy temperature is not enough to capture the peak temperatures and heat stress experienced by individual leaves in diverse tropical forest canopies. They also suggest that tropical trees have limited abilities to thermally acclimate to increasing temperatures.</p> <p><em>Keywords:</em> Canopy temperature, elevation gradient, fluorescence, heat tolerance, leaf area, leaf temperature, stomatal conductance, thermoregulation, tropical forest.</p>
<p>The responses of tropical forests to climate change depends on the ability of trees to acclimate to warming, as well as how interspecific variation in these responses affect tree community composition. In a unique tropical elevation gradient experiment in Rwanda, Rwanda TREE, we examine the sensitivity of tropical trees and forest stands to warming and altered water supply. Mixed multi-species plantations (20 tree species, 1800 trees per site) have been established at three sites with large variation in elevation (1300-2400 m) and climate (17-24 &#176;C mean daytime temperature), with additional water and nutrient manipulation treatments being applied at each site. Here we present an overview of results obtained this far regarding: (1) leaf gas exchange physiology; (2) photosynthetic heat tolerance; (3) water-use traits; (4) tree growth and mortality; (5) stand-level tree community composition. We also discuss the potential implications of our findings for the biodiversity and carbon storage of tropical forests in a changing climate.</p>
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