Scaling of petiole anatomies, mechanics and vasculatures with leaf size in the widespread Neotropical pioneer tree species Cecropia obtusa Trécul (Urticaceae).

Although the leaf economic spectrum has deepened our understanding of leaf trait variability, little is known about how leaf traits scale with leaf area. This uncertainty has resulted in the assumption that leaf traits should vary by keeping the same pace of variation with increases in leaf area across the leaf size range. We evaluated the scaling of morphological, tissue-surface and vascular traits with overall leaf area, and the functional significance of such scaling. We examined 1,271 leaves for morphological traits, and 124 leaves for anatomical and hydraulic traits, from 38 trees of Cecropia obtusa Trécul (Urticaceae) in French Guiana. Cecropia is a Neotropical genus of pioneer trees that can exhibit large laminas (0.4 m2 for C. obtusa), with leaf size ranging by two orders of magnitude. We measured (i) tissue fractions within petioles and their second moment of area, (ii) theoretical xylem hydraulic efficiency of petioles and (iii) the extent of leaf vessel widening within the hydraulic path. We found that different scaling of morphological trait variability allows for optimisation of lamina display among larger leaves, especially the positive allometric relationship between lamina area and petiole cross-sectional area. Increasing the fraction of pith is a key factor that increases the geometrical effect of supportive tissues on mechanical rigidity and thereby increases carbon-use efficiency. We found that increasing xylem hydraulic efficiency with vessel size results in lower leaf lamina area: xylem ratios, which also results in potential carbon savings for large leaves. We found that the vessel widening is consistent with hydraulic optimisation models. Leaf size variability modifies scaling of leaf traits in this large-leaved species.

Disentangling the effects of environment and ontogeny on tree functional dimensions for congeneric species in tropical forests.

Soil water and nutrient availability are key drivers of tree species distribution and forest ecosystem functioning, with strong species differences in water and nutrient use. Despite growing evidence for intraspecific trait differences, it remains unclear under which circumstances the effects of environmental gradients trump those of ontogeny and taxonomy on important functional dimensions related to resource use, particularly in tropical forests. Here, we explore how physiological, chemical, and morphological traits related to resource use vary between life stages in four species within the genus Micropholis that is widespread in lowland Amazonia. Specifically, we evaluate how environment, developmental stage, and taxonomy contribute to single-trait variation and multidimensional functional strategies. We find that environment, developmental stage, and taxonomy differentially contribute to functional dimensions. Habitats and seasons shape physiological and chemical traits related to water and nutrient use, whereas developmental stage and taxonomic identity impact morphological traits -especially those related to the leaf economics spectrum. Our findings suggest that combining environment, ontogeny, and taxonomy allows for a better understanding of important functional dimensions in tropical trees and highlights the need for integrating tree physiological and chemical traits with classically used morphological traits to improve predictions of tropical forests' responses to environmental change.

The relative weight of ontogeny, topology and climate in the architectural development of three North American conifers.

Knowledge of plant architecture allows retrospective study of plant development, hence provides powerful tools, through modelling and simulation, to link this development with environmental constraints, and then predict its response to global change. The present study aims to determine some of the main endogenous and exogenous variables driving the architectural development of three North American conifers. We measured architectural traits retrospectively on the trunk, branches and twigs of whole tree crowns for each species: annual shoot length (ASL), needle length, branching patterns and reproduction organs (male and female). We fitted a partial least square (PLS) regression to explain each architectural trait with respect to topological, ontogenic and climatic variables. Results showed a significant weight of these three groups of variables for previous and current year, corresponding, respectively, to organogenesis and elongation. Topological and ontogenic variables had the greatest weight in models. Particularly, all architectural traits were strongly correlated with ASL. We highlighted a negative architectural response of two species to higher than average temperatures, whereas the third one took advantage of these higher temperatures to some degree. Tree architectural development weekly but significantly improved with higher precipitation. Our study underlines the strong weight of topology and ontogeny in tree growth patterns at twig and branch scales. The correlation between ASL and other tree architectural traits should be integrated into architectural development models. Climate variables are secondary in importance at the twig scale. However, interannual climate variations influence all axis categories and branching orders and therefore significantly impact crown development as a whole. This latter impact may increase with climate change, especially as climate affects architectural traits over at least 2 years, through organogenesis and elongation.

Identifying source and formation altitudes of nitrates in drinking water from Réunion Island, France, using a multi-isotopic approach.

Nitrate concentrations, water isotopes (δ(2)H and δ(18)O(water)) and associated nitrate isotopes (δ(15)N(nitrate) and δ(18)O(nitrate)) from 10 drinking water wells, 5 fresh water springs and the discharge from 3 wastewater treatment stations in Réunion Island, located in the Indian Ocean, were analysed. We used a multi isotopic approach to investigate the extent of nitrate contamination, nitrate formation altitude and source of nitrates in Réunion Island's principal aquifer. Water from these study sites contained between 0.1 and 85.3 mg/L nitrate. δ(15)N(nitrate) values between +6 and +14‰ suggested the main sources of contamination were animal and/or human waste, rather than inorganic (synthetic) fertilisers, infiltrating through the subsurface into the saturated zone, due to rainfall leaching of the unsaturated zone at various altitudes of precipitation. Based on δ(15)N(nitrate) values alone, it was not possible to distinguish between animal and human activities responsible for the contamination of each specific catchment. However, using a multi isotope approach (δ(18)O(water) and δ(15)N(nitrate)), it was possible to relate the average altitude of rainfall infiltration (δ(18)O(water)) associated with the nitrate contamination (δ(18)O(nitrate)). This relationship between land use, rainfall recharge altitude and isotopic composition (δ(15)N(nitrate) and δ(18)O(water)) discriminated between the influences of human waste at lower (below 600 m elevation) or animal derived contamination (at elevations between 600 and 1300 m). By further comparing the theoretical altitude of nitrate formation calculated by the δ(18)O(nitrate), it was possible to determine that only 5 out of 15 fresh water wells and springs followed the conservative nitrate formation mechanism of 2/3δ(18)O(water)+1/3δ(18)O(air), to give nitrate formation altitudes which corresponded to land use activities.

The genus Cecropia: a biological clock to estimate the age of recently disturbed areas in the Neotropics.

Forest successional processes following disturbance take decades to play out, even in tropical forests. Nonetheless, records of vegetation change in this ecosystem are scarce, increasing the importance of the chronosequence approach to study forest recovery. However, this approach requires accurate dating of secondary forests, which until now was a difficult and/or expensive task. Cecropia is a widespread and abundant pioneer tree genus of the Neotropics. Here we propose and validate a rapid and straightforward method to estimate the age of secondary forest patches based on morphological observations of Cecropia trees. We found that Cecropia-inferred ages were highly correlated with known ages of the forest. We also demonstrate that Cecropia can be used to accurately date disturbances and propose twenty-one species distributed all over the geographical range of the genus as potential secondary forest chronometer species. Our method is limited in applicability by the maximal longevity of Cecropia individuals. Although the oldest chronosequence used in this study was 20 years old, we argue that at least for the first four decades after disturbance, the method described in this study provides very accurate estimations of secondary forest ages. The age of pioneer trees provides not only information needed to calculate the recovery of carbon stocks that would help to improve forest management, but also provides information needed to characterize the initial floristic composition and the rates of species remigration into secondary forest. Our contribution shows how successional studies can be reliably and inexpensively extended without the need to obtain forest ages based on expensive or potentially inaccurate data across the Neotropics.

Hydraulic architecture correlates with bud organogenesis and primary shoot growth in beech (Fagus sylvatica).

In beech (Fagus sylvatica L.), the number of leaf primordia preformed in the buds determines the length and the type (long versus short) of annual growth units, and thus, branch growth and architecture. We analyzed the correlation between the number of leaf primordia and the hydraulic conductance of the vascular system connected to the buds. Terminal buds of short growth units and axillary buds of long growth units on lower branches of mature trees were examined. Buds with less than four and more than five leaf primordia formed short and long growth units, respectively. Irrespective of the type of growth unit the bud was formed on, the occurrence of a large number of leaf primordia was associated with high xylem hydraulic conductance. Xylem conductance was correlated to the area of the outermost annual ring. These results suggest that organogenesis and primary growth in buds correlates with secondary growth of the growth units and thus with their hydraulic architecture. Possible causal relationships between the variables are discussed.

Epicormic branches: a growth indicator for the tropical forest tree, Dicorynia guianensis Amshoff (Caesalpiniaceae).

Architectural analyses of temperate tree species using a chronological approach suggest that the expression of epicormic branches is closely related to low growth rates in the axes that make up the branching system. Therefore, sole consideration of epicormic criteria may be sufficient to identify trees with low secondary growth levels or with both low primary and secondary growth levels. In a tropical tree such as Dicorynia guianensis (basralocus), where chronological studies are difficult, this relationship could be very useful as an easily accessible indicator of growth potentials. A simple method of architectural tree description was used to characterize the global structure of more than 1650 basralocus trees and to evaluate their growth level. Measurements of simple growth characters [height, basal diameter, internode length of submittal part (top of the main axis of the tree)] and the observation of four structural binary descriptors on the main stem (presence of sequential branches and young epicormic branches, state of the submittal part, global orientation), indicated that epicormic branch formation is clearly related to a decrease in length of the successive growth units of the main stem. Analysis of height vs. diameter ratios among different tree subgroups, with and without epicormic branching, suggested that trees with epicormic branches generally have a low level of secondary growth compared with primary growth.

Developmental patterns of the invasive bramble (Rubus alceifolius Poiret, Rosaceae) in Réunion island: an architectural and morphometric analysis.

The aim of this study was to identify the developmental stages of Rubus alceifolius and to determine one or more characteristic morphological markers for each stage. The developmental reconstitution method used involved a detailed description of many individuals throughout the different stages of growth, from germination to the development of an adult shoot capable of fruiting. Results revealed that R. alceifolius passes through five developmental stages that can be distinguished by changes in several morphological markers such as internode length and diameter, pith diameter and plant shape. This analysis indicated that R. alceifolius has a heteroblastic developmental pattern, midway between that of a bush and a liana. Moreover, results showed that this species taps environmental resources early in its development, i.e. foliarization is high (the foliar component overrides the caulinary component) and an autotrophic stage is rapidly reached, whereas it 'explores' the environment during the adult stage, i.e. axialization is substantial (the caulinary component overrides the foliar component) and autotrophy occurs at a later stage. The morphological markers identified could benefit land-use managers attempting to control this species before it reaches its optimum developmental stage.