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Background: Plants allocate resources to growth, defense, and stress resistance, and resource availability can affect the balance between these allocations. Allocation patterns are well-known to differ among species, but what controls possible intra-specific trade-offs and if variation in growth vs. defense potentially evolves in adaptation to resource availability. Methods: We measured growth and defense in a provenance trial of rubber trees (Hevea brasiliensis) with clones originating from the Amazon basin. To test hypotheses on the allocation to growth vs. defense, we relate biomass growth and latex production to wood and leaf traits, to climate and soil variables from the location of origin, and to the genetic relatedness of the Hevea clones. Results: Contrary to expectations, there was no trade-off between growth and defense, but latex yield and biomass growth were positively correlated, and both increased with tree size. The absence of a trade-off may be attributed to the high resource availability in a plantation, allowing trees to allocate resources to both growth and defense. Growth was weakly correlated with leaf traits, such as leaf mass per area, intrinsic water use efficiency, and leaf nitrogen content, but the relative investment in growth vs. defense was not associated with specific traits or environmental variables. Wood and leaf traits showed clinal correlations to the rainfall and soil variables of the places of origin. These traits exhibited strong phylogenetic signals, highlighting the role of genetic factors in trait variation and adaptation. The study provides insights into the interplay between resource allocation, environmental adaptations, and genetic factors in trees. However, the underlying drivers for the high variation of latex production in one of the commercially most important tree species remains unexplained.

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The Bromeliaceae family has been used as a model to study adaptive radiation due to its terrestrial, epilithic, and epiphytic habits with wide morpho-physiological variation. Functional groups described by Pittendrigh in 1948 have been an integral part of ecophysiological studies. In the current study, we revisited the functional groups of epiphytic bromeliads using a 204 species trait database sampled throughout the Americas. Our objective was to define epiphytic functional groups within bromeliads based on unsupervised classification, including species from the dry to the wet end of the Neotropics. We performed a hierarchical cluster analysis with 16 functional traits and a discriminant analysis, to test for the separation between these groups. Herbarium records were used to map species distributions and to analyze the climate and ecosystems inhabited. The clustering supported five groups, C3 tank and CAM tank bromeliads with deep tanks, while the atmospheric group (according to Pittendrigh) was divided into nebulophytes, bromeliads with shallow tanks, and bromeliads with pseudobulbs. The two former groups showed distinct traits related to resource (water) acquisition, such as fog (nebulophytes) and dew (shallow tanks). We discuss how the functional traits relate to the ecosystems inhabited and the relevance of acknowledging the new functional groups.

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The reasons why the range size of closely related species often varies significantly have intrigued scientists for many years. Among other hypotheses, species with high trait variation were suggested to occupy more diverse environments, have more continuity in their distributions, and consequently have larger range sizes. Here, using 34 tree species of lowlands tropical rainforest in southern Costa Rica, we explored whether inherent trait variability expressed at the local scale in functional traits is related to the species' total geographical range size. We formed 17 congeneric pairs of one narrow endemic and one widespread species, sampled 335 individuals and measured eight functional traits: leaf area, leaf thickness, leaf dry matter content, specific leaf area, leaf nitrogen content, leaf phosphorus content, leaf nitrogen to phosphorus ratio, and wood specific gravity. We tested whether there are significant differences in the locally expressed variation of individual traits or in multidimensional trait variance between the species in congeneric pairs and whether species' range size could hence be predicted from local trait variability. However, we could not find such differences between widely distributed and narrow range species. We discuss the possible reasons for these findings including the fact that higher trait variability of widespread species may result from successive local adaptations during range expansion and may hence often be an effect rather than the cause of larger ranges.

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Wood structure differs widely among tree species and species with faster growth, higher mortality and larger maximum size have been reported to have fewer but larger vessels and higher hydraulic conductivity (Kh). However, previous studies compiled data from various sources, often failed to control tree size and rarely controlled variation in other traits. We measured wood density, tree size and vessel traits for 325 species from a wet forest in Panama, and compared wood and leaf traits to demographic traits using species-level data and phylogenetically independent contrasts. Wood traits showed strong phylogenetic signal whereas pairwise relationships between traits were mostly phylogenetically independent. Trees with larger vessels had a lower fraction of the cross-sectional area occupied by vessel lumina, suggesting that the hydraulic efficiency of large vessels permits trees to dedicate a larger proportion of the wood to functions other than water transport. Vessel traits were more strongly correlated with the size of individual trees than with maximal size of a species. When individual tree size was included in models, Kh scaled positively with maximal size and was the best predictor for both diameter and biomass growth rates, but was unrelated to mortality.

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Anthropogenic nitrogen deposition is currently causing a more than twofold increase of reactive nitrogen input over large areas in the tropics. Elevated (15)N abundance (δ(15)N) in the growth rings of some tropical trees has been hypothesized to reflect an increased leaching of (15)N-depleted nitrate from the soil, following anthropogenic nitrogen deposition over the last decades. To find further evidence for altered nitrogen cycling in tropical forests, we measured long-term δ(15)N values in trees from Bolivia, Cameroon, and Thailand. We used two different sampling methods. In the first, wood samples were taken in a conventional way: from the pith to the bark across the stem of 28 large trees (the "radial" method). In the second, δ(15)N values were compared across a fixed diameter (the "fixed-diameter" method). We sampled 400 trees that differed widely in size, but measured δ(15)N in the stem around the same diameter (20 cm dbh) in all trees. As a result, the growth rings formed around this diameter differed in age and allowed a comparison of δ(15)N values over time with an explicit control for potential size-effects on δ(15)N values. We found a significant increase of tree-ring δ(15)N across the stem radius of large trees from Bolivia and Cameroon, but no change in tree-ring δ(15)N values over time was found in any of the study sites when controlling for tree size. This suggests that radial trends of δ(15)N values within trees reflect tree ontogeny (size development). However, for the trees from Cameroon and Thailand, a low statistical power in the fixed-diameter method prevents to conclude this with high certainty. For the trees from Bolivia, statistical power in the fixed-diameter method was high, showing that the temporal trend in tree-ring δ(15)N values in the radial method is primarily caused by tree ontogeny and unlikely by a change in nitrogen cycling. We therefore stress to account for tree size before tree-ring δ(15)N values can be properly interpreted.

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[1] Natural archives of oxygen isotopes in precipitation may be used to study changes in the hydrological cycle in the tropics, but their interpretation is not straightforward. We studied to which degree tree rings of Mimosa acantholoba from southern Mexico record variation in isotopic composition of precipitation and which climatic processes influence oxygen isotopes in tree rings (δ18Otr). Interannual variation in δ18Otr was highly synchronized between trees and closely related to isotopic composition of rain measured at San Salvador, 710 km to the southwest. Correlations with δ13C, growth, or local climate variables (temperature, cloud cover, vapor pressure deficit (VPD)) were relatively low, indicating weak plant physiological influences. Interannual variation in δ18Otr correlated negatively with local rainfall amount and intensity. Correlations with the amount of precipitation extended along a 1000 km long stretch of the Pacific Central American coast, probably as a result of organized storm systems uniformly affecting rainfall in the region and its isotope signal; episodic heavy precipitation events, of which some are related to cyclones, deposit strongly 18O-depleted rain in the region and seem to have affected the δ18Otr signal. Large-scale controls on the isotope signature include variation in sea surface temperatures of tropical north Atlantic and Pacific Ocean. In conclusion, we show that δ18Otr of M. acantholoba can be used as a proxy for source water δ18O and that interannual variation in δ18Oprec is caused by a regional amount effect. This contrasts with δ18O signatures at continental sites where cumulative rainout processes dominate and thus provide a proxy for precipitation integrated over a much larger scale. Our results confirm that processes influencing climate-isotope relations differ between sites located, e.g., in the western Amazon versus coastal Mexico, and that tree ring isotope records can help in disentangling the processes influencing precipitation δ18O.

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Deposition of reactive nitrogen (N) from human activities has large effects on temperate forests where low natural N availability limits productivity but is not known to affect tropical forests where natural N availability is often much greater. Leaf N and the ratio of N isotopes (δ(15)N) increased substantially in a moist forest in Panama between ~1968 and 2007, as did tree-ring δ(15)N in a dry forest in Thailand over the past century. A decade of fertilization of a nearby Panamanian forest with N caused similar increases in leaf N and δ(15)N. Therefore, our results indicate regional increases in N availability due to anthropogenic N deposition. Atmospheric nitrogen dioxide measurements and increased emissions of anthropogenic reactive N over tropical land areas suggest that these changes are widespread in tropical forests.

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Direct or indirect anthropogenic effects on ecosystem nitrogen cycles are important components of global change. Recent research has shown that N isotopes in tree rings reflect changes in ecosystem nitrogen sources or cycles and can be used to study past changes. We analyzed trends in two tree species from a remote and pristine tropical rainforest in Brazil, using trees of different ages to distinguish between the effect of tree age and long-term trends. Because sapwood differed from heartwood in delta(15)N and N concentration and N can be translocated between living sapwood cells, long-term trends are best seen in dead heartwood. Heartwood delta(15)N in Spanish cedar (Cedrela odorata) and big-leaf mahogany (Swietenia macrophylla) increased with tree age, and N concentrations increased with age in Cedrela. Controlling for tree age, delta(15)N increased significantly during the past century even when analyzing only heartwood and after removing labile N compounds. In contrast to northern temperate and boreal forests where wood delta(15)N often decreased, the delta(15)N increase in a remote rainforest is unlikely to be a direct signal of changed N deposition. More plausibly, the change in N isotopic composition indicates a more open N cycle, i.e., higher N losses relative to internal N cycling in the forest, which could be the result of changed forest dynamics.

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Elevated CO(2) concentrations ([CO(2)]) affect plant water relations and photosynthesis, and the increase in atmospheric [CO(2)] over the past 100-200 years has been related to changes in stomatal density and the carbon isotope ratio (delta(13)C) in tree rings and leaves from herbarium specimens. Because many tropical trees do not produce annual growth rings and their wood is therefore difficult to date, no trends in delta(13)C of tropical trees have been reported. Wood from Cedrela odorata L. (tropical cedar) and Swietenia macrophylla King (bigleaf mahogany), which do produce annual rings, was collected from a primary rain forest in Aripuanã, Brazil (10 degrees 09' S, 59 degrees 26' W). We measured wood cellulose delta(13)C in 10-year growth increments from 37 Cedrela trees (between 11 and 151 years old in 2001) and 16 Swietenia trees (48-126 years old). A comparison of delta(13)C in cellulose of trees from different decades and of trees of different cambial ages showed that the amount of delta(13)C was largely related to the decade the wood was produced in, and not, or only to a minor extent, to tree age. Cellulose delta(13)C decreased from -26.0 to -27.3 per thousand in Cedrela and from -25.7 to -27.1 per thousand in Swietenia, with the largest changes occurring during the past 50 years. Based on these data and the trends in atmospheric [CO(2)] and delta(13)CO(2), we calculated that the internal [CO(2)] increased from about 220 to 260 ppm and that intrinsic water-use efficiency increased by 34% in Cedrela and by 52% in Swietenia. This may have implications for the water cycle and may explain the trend toward increased tree growth and turnover observed in some tropical forests.

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BACKGROUND AND AIMS: Seeds of epiphytes must land on branches with suitable substrates and microclimates to germinate and for the resulting seedlings to survive. It is important to understand the fate of seeds and seedlings in order to model populations, but this is often neglected when only established plants are included in analyses. METHODS: The seeds of five bromeliad species were exposed to different canopy positions in a Mexican montane forest, and germination and early seedling survival were recorded. Additionally, the survival of naturally dispersed seedlings was monitored in a census over 2.5 years. Survival analysis, a procedure rarely used in plant ecology, was used to study the influence of branch characteristics and light on germination and seedling survival in natural and experimental populations. KEY RESULTS: Experimental germination percentages ranged from 7.2 % in Tillandsia deppeana to 33.7 % in T. juncea, but the seeds of T. multicaulis largely failed to germinate. Twenty months after exposure between 3.5 and 9.4 % of the seedlings were still alive. There was no evidence that canopy position affected the probability of germination, but time to germination was shorter in less exposed canopy positions indicating that higher humidity accelerates germination. More experimental seedlings survived when canopy openness was high, whereas survival in census-seedlings was influenced by moss cover. While mortality decreased steadily with age in juveniles of the atmospheric Tillandsia, in the more mesomorphic Catopsis sessiliflora mortality increased dramatically in the dry season. CONCLUSIONS: Seedling mortality, rather than the failure to germinate, accounts for the differential distribution of epiphytes within the canopy studied. With few safe sites to germinate and high seedling mortality, changes of local climate may affect epiphyte populations primarily through their seedling stage.

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