Plasticity and implications of water-use traits in contrasting tropical tree species under climate change.

Plants face a trade-off between hydraulic safety and growth, leading to a range of water-use strategies in different species. However, little is known about such strategies in tropical trees and whether different water-use traits can acclimate to warming. We studied five water-use traits in 20 tropical tree species grown at three different altitudes in Rwanda (RwandaTREE): stomatal conductance (gs), leaf minimum conductance (gmin), plant hydraulic conductance (Kplant), leaf osmotic potential (ψo) and net defoliation during drought. We also explored the links between these traits and growth and mortality data. Late successional (LS) species had low Kplant, gs and gmin and, thus, low water loss, while low ψo helped improve leaf water status during drought. Early successional (ES) species, on the contrary, used more water during both moist and dry conditions and exhibited pronounced drought defoliation. The ES strategy was associated with lower mortality and more pronounced growth enhancement at the warmer sites compared to LS species. While Kplant and gmin showed downward acclimation in warmer climates, ψo did not acclimate and gs measured at prevailing temperature did not change. Due to distinctly different water use strategies between successional groups, ES species may be better equipped for a warmer climate as long as defoliation can bridge drought periods.

Contrasting warming responses of photosynthesis in early- and late-successional tropical trees.

The productivity and climate feedbacks of tropical forests depend on tree physiological responses to warmer and, over large areas, seasonally drier conditions. However, knowledge regarding such responses is limited due to data scarcity. We studied the impact of growth temperature on net photosynthesis (An), maximum rates of Rubisco carboxylation at 25 °C (Vcmax25), stomatal conductance (gs) and the slope parameter of the stomatal conductance-photosynthesis model (g1), in 10 early successional (ES) and 8 late-successional (LS) tropical tree species grown at three sites along an elevation gradient in Rwanda, differing by 6.8 °C in daytime ambient air temperature. The effect of seasonal drought on An was also investigated. We found that warm climate decreased wet-season An in LS species, but not in ES species. Values of Vcmax25 were lower at the warmest site across both successional groups, and An and Vcmax25 were higher in ES compared with LS species. Stomatal conductance exhibited no significant site differences and g1 was similar across both sites and successional groups. Drought strongly reduced An at warmer sites but not at the coolest montane site and this response was similar in both ES and LS species. Our results suggest that warming has negative effects on leaf-level photosynthesis in LS species, while both LS and ES species suffer photosynthesis declines in a warmer climate with more pronounced droughts. The contrasting responses of An between successional groups may lead to shifts in species' competitive balance in a warmer world, to the disadvantage of LS trees.

Handling the heat - photosynthetic thermal stress in 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 (Tleaf ) across the species. At the warmest sites, the thermal threshold for normal function of photosystem II was exceeded in the species with the highest Tleaf despite their higher heat tolerance. This was not the case in the species with the highest transpiration rates and lowest Tleaf . 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.

Limited thermal acclimation of photosynthesis in tropical montane tree species.

The temperature sensitivity of physiological processes and growth of tropical trees remains a key uncertainty in predicting how tropical forests will adjust to future climates. In particular, our knowledge regarding warming responses of photosynthesis, and its underlying biochemical mechanisms, is very limited. We grew seedlings of two tropical montane rainforest tree species, the early-successional species Harungana montana and the late-successional species Syzygium guineense, at three different sites along an elevation gradient, differing by 6.8℃ in daytime ambient air temperature. Their physiological and growth performance was investigated at each site. The optimum temperature of net photosynthesis (ToptA ) did not significantly increase in warm-grown trees in either species. Similarly, the thermal optima (ToptV and ToptJ ) and activation energies (EaV and EaJ ) of maximum Rubisco carboxylation capacity (Vcmax ) and maximum electron transport rate (Jmax ) were largely unaffected by warming. However, Vcmax , Jmax and foliar dark respiration (Rd ) at 25℃ were significantly reduced by warming in both species, and this decline was partly associated with concomitant reduction in total leaf nitrogen content. The ratio of Jmax /Vcmax decreased with increasing leaf temperature for both species, but the ratio at 25℃ was constant across sites. Furthermore, in H. montana, stomatal conductance at 25℃ remained constant across the different temperature treatments, while in S. guineense it increased with warming. Total dry biomass increased with warming in H. montana but remained constant in S. guineense. The biomass allocated to roots, stem and leaves was not affected by warming in H. montana, whereas the biomass allocated to roots significantly increased in S. guineense. Overall, our findings show that in these two tropical montane rainforest tree species, the capacity to acclimate the thermal optimum of photosynthesis is limited while warming-induced reductions in respiration and photosynthetic capacity rates are tightly coupled and linked to responses of leaf nitrogen.

Complete or overcompensatory thermal acclimation of leaf dark respiration in African tropical trees.

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 (Rd ), to these new conditions. However, knowledge on this is currently limited due to data scarcity. We studied the impact of growth temperature on Rd and its dependency on net photosynthesis (An ), leaf nitrogen (N) and phosphorus (P) contents, and leaf mass per unit area (LMA) in 16 early-successional (ES) and late-successional (LS) tropical tree species in multispecies plantations along an elevation gradient (Rwanda TREE project). Moreover, we explored the effect of drought on Rd in one ES and one LS species. Leaf Rd at 20°C decreased at warmer sites, regardless if it was expressed per unit leaf area, mass, N or P. This acclimation resulted in an 8% and a 28% decrease in Rd at prevailing nighttime temperatures in trees at the intermediate and warmest sites, respectively. Moreover, drought reduced Rd , particularly in the ES species and at the coolest site. Thermal acclimation of Rd is complete or overcompensatory and independent of changes in leaf nutrients or LMA in African tropical trees.

Urticaceae leaves with stinging trichomes were already present in latest early Eocene Okanogan Highlands, British Columbia, Canada.

PREMISE: Paleontologists use tooth form to assess diets of fossil mammals. Plants would also be expected to adapt their morphology to respond to herbivory. Fossil nettle leaves with definitive stinging trichomes (tribe Urticeae, family Urticaceae) are described from the early Eocene upland lacustrine floras of the Okanogan Highlands, British Columbia, Canada. This is the first report of stinging trichomes in the fossil record. Their occurrence in western North America at a time of major large herbivorous mammal radiation suggests they acted, as they do today, as a deterrent for mammal herbivory. METHODS: Fossil leaf compressions and extant leaves were photographed with standard methods. Focus-shift stacking was used to layer photos of the fossil leaves. RESULTS: Urticaceous fossil leaves from the Okanogan Highlands greatly resemble their modern relatives in leaf morphology and particularly in both stinging and nonstinging trichomes. Nettles are common components of the flora of the Volcanoes National Park in Rwanda. This region is used as a modern analogue for the Okanogan Highlands, based on comparable elevation, equable conditions that host both similar floras and large folivores. CONCLUSIONS: Nettles in tribe Urticeae (Urticaceae) producing leaves with stinging and nonstinging trichomes were already present in the early Eocene of western North America at a pivotal time during the early radiation of modern mammalian herbivore groups. They offer tantalizing evidence of a selective response that plants may have developed to protect themselves from the evolving mammalian herbivores of that time.