The effect of temperature change on leaf physiology has been extensively studied in temperate trees and to some extent in boreal and tropical tree species. While increased temperature typically stimulates leaf CO2 assimilation and tree growth in high-altitude ecosystems, tropical species are often negatively affected. These trees may operate close to their temperature optima and have a limited thermal acclimation capacity due to low seasonal and historical variation in temperature. To test this hypothesis, we studied the extent to which the temperature sensitivities of leaf photosynthesis and respiration acclimate to growth temperature in four common African tropical tree species. Tree seedlings native to different altitudes and therefore adapted to different growth temperatures were cultivated at three different temperatures in climate-controlled chambers. We estimated the acclimation capacity of the temperature sensitivities of light-saturated net photosynthesis, the maximum rates of Rubisco carboxylation (Vcmax) and thylakoid electron transport (J), and dark respiration. Leaf thylakoid membrane lipid composition, nitrogen content and leaf mass per area were also analyzed. Our results showed that photosynthesis in tropical tree species acclimated to higher growth temperatures, but that this was weakest in the species originating from the coolest climate. The temperature optimum of J acclimated significantly in three species and variation in J was linked to changes in the thylakoid membrane lipid composition. For Vcmax, there was only evidence of significant acclimation of optimal temperature in the lowest elevation species. Respiration acclimated to maintain homeostasis at growth temperature in all four species. Our results suggest that the lowest elevation species is better physiologically adapted to acclimate to high growth temperatures than the highest elevation species, indicating a potential shift in competitive balance and tree community composition to the disadvantage of montane tree species in a warmer world.
Tropical canopies are complex, with multiple canopy layers and pronounced gap dynamics contributing to their high species diversity and productivity. An important reason for this complexity is the large variation in shade tolerance among different tree species. At present, we lack a clear understanding of which plant traits control this variation, e.g., regarding the relative contributions of whole-plant versus leaf traits or structural versus physiological traits. We investigated a broad range of traits in six tropical montane rainforest tree species with different degrees of shade tolerance, grown under three different radiation regimes (under the open sky or beneath sparse or dense canopies). The two distinct shade-tolerant species had higher fractional biomass in leaves and branches while shade-intolerant species invested more into stems, and these differences were greater under low radiation. Leaf respiration and photosynthetic light compensation point did not vary with species shade tolerance, regardless of radiation regime. Leaf temperatures in open plots were markedly higher in shade-tolerant species due to their low transpiration rates and large leaf sizes. Our results suggest that interspecific variation in shade tolerance of tropical montane trees is controlled by species differences in whole-plant biomass allocation strategy rather than by difference in physiological leaf traits determining leaf carbon balance at low radiation.
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.
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 rain forests (TMFs) are considered particularly vulnerable to climate change, but this hypothesis remains poorly evaluated due to data scarcity. To reduce the knowledge gap in the response of TMF trees to warming, we established a field experiment along a 1300–2400 m elevation gradient as a proxy for warming in Rwanda. Seedling-size trees of 20 species native to montane forests in eastern and central Africa were planted in multi-species plots at three sites along the gradient. They have overlapping distributions but primarily occur in either transitional rain forests (∼ 1600–2000 ) or mid-elevation TMFs (∼ 2000–3000 ), 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 2 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 rain forest origin strongly increased proportional to 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 of becoming outcompeted by lower-elevation and ES species due to a thermophilisation response in a warmer climate has important implications for biodiversity and carbon storage of Afromontane forests.
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.