Previous studies of heat tolerance of tropical trees have focused on canopy leaves exposed to full sunlight and high temperatures. However, in lowland tropical forests with leaf area indices of 5-6, the vast majority of leaves experience varying degrees of shade and a reduced heat load compared to sun leaves. Here we tested whether heat tolerance is lower in shade than in sun leaves. For three tropical tree species, Calophyllum inophyllum, Inga spectabilis, and Ormosia macrocalyx, disks of fully developed shade and sun leaves were subjected to 15-min heat treatments, followed by measurement of chlorophyll a fluorescence after 48 h of recovery. In two of the three species, the temperature causing a 50% decrease of the fluorescence ratio F/F (T) was significantly lower (by ~ 1.0 °C) in shade than in sun leaves, indicating a moderately decreased heat tolerance of shade leaves. In shade leaves of these two species, the rise in initial fluorescence, F, also occurred at lower temperatures. In the third species, there was no shade-sun difference in T. In situ measurements of photosynthetic CO assimilation showed that the optimum temperature for photosynthesis tended to be lower in shade leaves, although differences were not significant. At supra-optimal temperatures, photosynthesis was largely constrained by stomatal conductance, and the high-temperature CO compensation point, T, occurred at considerably lower temperatures than T. Our study demonstrates that the temperature response of shade leaves of tropical trees differs only marginally from that of sun leaves, both in terms of heat tolerance and photosynthetic performance.
More frequent droughts and rising temperatures pose serious threats to tropical forests. When stomata are closed under dry and hot conditions, plants lose water through leaf cuticles, but little is known about cuticle conductance (g min ) of tropical trees, how it varies among species and environments, and how it is affected by temperature. We determined g min in relation to temperature for 24 tropical tree species across a steep rainfall gradient in Panama, by recording leaf drying curves at different temperatures in the laboratory. In contrast to our hypotheses, g min did not differ systematically across the rainfall gradient; species differences did not reflect phylogenetic patterns; and in most species g min did not significantly increase between 25 and 50°C. g min was higher in deciduous than in evergreen species, in species with leaf trichomes than in species without, in sun leaves than in shade leaves, and tended to decrease with increasing leaf mass per area across species. There was no relationship between stomatal and cuticle conductance. Large species differences in g min and its temperature response suggest that more frequent hot droughts may lead to differential survival among tropical tree species, regardless of species' position on the rainfall gradient.
Almost 40 years ago, Terry L. Erwin published a seemingly audacious proposition: There may be as many as 30 million species of insects in the world. Here, we translate Erwin's verbal argument into a diversity‐ratio model—the Erwin Equation of Biodiversity—and discuss how it has inspired other biodiversity estimates. We categorize, describe the assumptions for, and summarize the most commonly used methods for calculating estimates of global biodiversity. Subsequent diversity‐ratio extrapolations have incorporated parameters representing empirical insect specialization ratios, and how insect specialization changes at different spatial scales. Other approaches include macroecological diversity models and diversity curves. For many insect groups with poorly known taxonomies, diversity estimates are based on the opinions of taxonomic experts. We illustrate our current understanding of insect diversity by focusing on the six most speciose insect orders worldwide. For each order, we compiled estimates of the (a) maximum estimated number of species, (b) minimum estimated number of species, and (c) number of currently described species. By integrating these approaches and considering new information, we believe an estimate of 5.5 million species of insects in the world is much too low. New molecular methodologies (e.g., metabarcoding and NGS studies) are revealing daunting numbers of cryptic and previously undescribed species, at the same time increasing our precision but also uncertainty about present estimates. Not until technologies advance and sampling become more comprehensive, especially of tropical biotas, will we be able to make robust estimates of the total number of insect species on Earth.Abstract in Spanish is available with online material.
Photosynthetic carbon uptake by tropical forests is of critical importance in regulating the earth’s climate, but rising temperatures threaten this stabilizing influence of tropical forests. Most research on how temperature affects photosynthesis focuses on fully sun-exposed leaves, and little is known about shade leaves, even though shade leaves greatly outnumber sun leaves in lowland tropical forests. We measured temperature responses of light-saturated photosynthesis, stomatal conductance, and the biochemical parameters VCMax (maximum rate of RuBP carboxylation) and JMax (maximum rate of RuBP regeneration, or electron transport) on sun and shade leaves of mature tropical trees of three species in Panama. As expected, biochemical capacities and stomatal conductance were much lower in shade than in sun leaves, leading to lower net photosynthesis rates. However, the key temperature response traits of these parameters—the optimum temperature (TOpt) and the activation energy—did not differ systematically between sun and shade leaves. Consistency in the JMax to VCMax ratio further suggested that shade leaves are not acclimated to lower temperatures. For both sun and shade leaves, stomatal conductance had the lowest temperature optimum (~25 °C), followed by net photosynthesis (~30 °C), JMax (~34 °C) and VCMax (~38 °C). Stomatal conductance of sun leaves decreased more strongly with increasing vapor pressure deficit than that of shade leaves. Consistent with this, modeled stomatal limitation of photosynthesis increased with increasing temperature in sun but not shade leaves. Collectively, these results suggest that modeling photosynthetic carbon uptake in multi-layered canopies does not require independent parameterization of the temperature responses of the biochemical controls over photosynthesis of sun and shade leaves. Nonetheless, to improve the representation of the shade fraction of carbon uptake dynamics in tropical forests, better understanding of stomatal sensitivity of shade leaves to temperature and vapor pressure deficit will be required.
1. Tropical wet forest plants experience relatively stable temperatures throughout the year. However, tropical forests represent a mosaic of habitats characterized by different temperatures. Heat tolerances are expected to be adapted to temperatures specific to their habitats. Although heat tolerance of species sharing similar environments is expected to be similar, it is also possible that heat tolerance is constrained by evolutionary history because closely related species usually display similar physiologies. When exotic species are introduced to novel communities, colonization may be facilitated by their previous adaptation to high temperatures and other physiological, genetic and demographic traits, which may grant them some competitive advantage. Increasing temperatures may represent a strong environmental filter affecting community assembly, and higher heat tolerances could facilitate the persistence of exotic species in novel environments.2. Using a community of 32 native and 7 exotic Zingiberales species from different tropical habitats in Costa Rica, Central America, we aim to answer the following questions: (a) does evolutionary history constrain heat tolerance? (b) do plants in the same habitat display similar heat tolerances? (c) do the heat tolerances of exotic species differ from those of native species? 3. We measured temperature-dependent changes in photosynthetic fluorescence to determine the temperature at which the first sign of damage to photosystem II is observed (T 15 ), and the temperature at which the fluorescence of photosystem II is reduced by 50% (T 50 ). Using a community phylogeny, we tested for phylogenetic signal in T 15 and T 50 . In addition, we tested for differences in heat tolerance among Zingiberales from old growth, secondary forests and open areas, as well as between native and exotic species.
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