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Variation in Leaf Morphology and Branching Pattern of Some Tropical Rain Forest Species from Guadeloupe

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Variation in Leaf Morphology and Branching Pattern of Some Tropical Rain Forest Species from Guadeloupe Variation in leaf morphology and branching pattern of some tropical rain forest species from Guadeloupe (French West Indies) under semi-controlled light conditions M Ducrey To cite this version: M Ducrey. Variation in leaf morphology and branching pattern of some tropical rain forest species from Guadeloupe (French West Indies) under semi-controlled light conditions. Annales des sciences forestières, INRA/EDP Sciences, 1992, 49 (6), pp.553-570. hal-00882820 HAL Id: hal-00882820 https://hal.archives-ouvertes.fr/hal-00882820 Submitted on 1 Jan 1992 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Original article Variation in leaf morphology and branching pattern of some tropical rain forest species from Guadeloupe (French West Indies) under semi-controlled light conditions M Ducrey INRA, Laboratoire de Recherches Forestières Méditerranéennes, Avenue A Vivaldi, F-84000 Avignon, France (Received 18 March 1992; accepted 7 July 1992) Summary — Seedlings of 7 canopy species from the Guadeloupe tropical rain forest (Dacryodes excelsa, Amanoa caribaea, Richeria grandis, Simaruba amara, Symphonia globulifera, Byrsonima coriacea and Podocarpus coriaceus) were raised in full sunlight and under artifical neutral shade transmitting 6, 11, 19 and 54% light for 2 to 3 years. At the end of this period, the number of leaves and branches, leaf size, specific leaf area and stomatal density were observed for each plant. For all species, the maximum number of leaves was obtained in partial shade (11 or 19% sunlight). Branch- ing occurrence depended more on species type than on light conditions. Both individual leaf size and specific leaf area increased regularly with shade, but in a proportion which varied according to the species. Stomatal density was highly variable from one species to another and increased with greater light. The morphological plasticity of species response to light conditions was then analysed and related to shade tolerance. In order of decreasing plasticity, the first species found were R gran- dis, S amara and B coriacea, which were the most plastic and the most shade intolerant, followed by A caribaea and P coriaceus, less plastic but shade-tolerant species. Finally, D excelsa and S globu- lifera were found to be the least plastic species and highly or moderately shade-tolerant. tropical rain forest / leaf morphology / specific leaf area / branching pattern / shade tolerance Résumé — Variations de la morphologie foliaire et branchaison de quelques espèces de la forêt tropicale humide de Guadeloupe en conditions semi-contrôlées d’éclairement. De jeunes semis de 7 espèces de la strate arborescente de la forêt tropicale humide de Guadeloupe (Da- cryodes excelsa, Amanoa caribaea, Richeria grandis, Simaruba amara, Symphonia globulifera, Byr- sonima coriacea et Podocarpus coriaceus) ont été élevés pendant 2-3 ans en pleine lumière et sous ombrages artificiels neutres laissant passer 6%, 11%, 19% et 54% de la pleine lumière. À la fin de cette période on a observé sur chaque plant, le nombre de feuilles et de ramifications, la taille et la surface spécifique des feuilles ainsi que la densité stomatique. Pour toutes les espèces étudiées, le nombre de feuilles est maximal pour des ombrages moyens (11 ou 19% de la pleine lumière). La présence de ramifications dépend davantage des espèces que des conditions d’éclairement. La sur- face individuelle des feuilles ainsi que leur surface spécifique augmentent régulièrement avec l’om- brage mais dans des proportions variables selon les espèces. La densité stomatique, très variable d’une espèce à l’autre, augmente avec l’éclairement. La plasticité morphologique des espèces en ré- ponse aux conditions d’éclairement est ensuite analysée et interprétée en termes de tolérance à l’om- brage. Par ordre de plasticité décroissante, on trouve R grandis, S amara et B coriacea qui sont les espèces les plus plastiques et les plus intolérantes à l’ombrage. On trouve ensuite A caribaea et P coriaceus, moins plastiques mais tolérantes à l’ombrage. D excelsa, et S globulifera sont les moins plastiques et sont modérément ou fortement tolérantes à l’ombrage. forêt tropicale humide / morphologie foliaire / surface foliaire spécifique / blanchaison / tolé- rance à l’ombrage INTRODUCTION studied under 2 different thinning intensi- ties. The variations in environmental condi- tions due to the different silvicultural treat- The reaction of trees to envi- varying light ments were then used as a means of ronments, particularly to shade, can be determining the range of requirements at different levels. First of at light compared all, in the species studied, from the most the we find which species level, species shade-intolerant to the most shade-tolerant. require full sunlight and others which are A silvicultural is not more or less shade-tolerant. On the indi- uniquely approach sufficient to understand the forest behavi- vidual level, within the same species or our of a and its relative genotype, we find trees which have grown given species in different light environments and have place in a forest succession. It therefore different phenotypes (shade phenotypes seemed of interest to further the know- on these mor- or sun phenotypes). Finally, within the ledge species by studying variations in leaves and branch- same individual, particularly within a stand, phological in response to light conditions sun and shade leaves are found, depend- ing pattern This is of value ing on their position in the tree crown. during growth. approach for 2 reasons. First of all, the use of mor- These facts are generally known for phological criteria to account for physiolog- most tree species growing in temperate cli- ical potentials under varying light condi- but have been less studied for mates, trop- tions appears to be possible using existing ical species. In particular, the shade re- relationships between physiological and of the main commercial in sponse species morphogenetic processes (Tsel’Niker, the rainforest of is tropical Guadeloupe 1977). Secondly, the range of morphologi- unknown. practically cal variations in the leaf system under ex- The experiments conducted (Ducrey, treme light conditions is a good means of 1982; Ducrey and Labbé, 1985) on stimu- determining the forest behaviour of a given lated and controlled natural regeneration species (Smith, 1982; Fetcher et al, 1983; in the Guadeloupe rainforest provided the Goulet and Bellefleur, 1986). first results (Ducrey and Labbé, 1986) on This article examines the morphological the forest behaviour of the main tree spe- variations in leaves and branching pattern cies favoured for natural regeneration. for 7 evergreen species subjected to 5 dif- Methods similar to the progressive felling ferent light conditions. The experiment also regeneration and the tropical shelterwood took into account photosynthetic response, system were adopted. Survival and growth growth and biomass production, which will of seedlings from different species were be discussed in further papers. MATERIALS AND METHODS full sunlight. The 4 tunnel shelters were 15 m long and 6 m wide and covered with reinforced transparent PVC as a protection against rainfall. Three of them were shaded with different black of Description seedlings neutral shade screens in order to obtain various of species studied shade conditions. Finally, global radiation meas- urements with Li-Cor pyranometers indicated 6.4% light under tunnel I, 11.4% under tunnel II, The used for the were seedlings experiment 18.8% under tunnel III and 54.3% under tunnel sampled from the tropical rainforest of Guade- IV. loupe, French West Indies. They came from the Table I summarizes climatic data under tun- "Débauchée" area (Ducrey, 1986) at an eleva- nel shelters. These were and oriented in tion of 250 m. Mean temperatures were 23 °C in opened January and 26 °C in July. Mean annual rainfall the direction of prevailing winds. The microcli- matic conditions under the tunnels were the was > 3 000 mm. There was a short dry season from January to April, but the monthly rainfall same as those in the open air treatment (meteo- data measured a weather was always > 100 mm. rological by station), except for tunnel IV whose maximum tempera- The 7 studied were domi- species evergreen tures were slightly higher than the others. This nant and co-dominant trees from the middle and could be explained, as the shade under this tun- late successional of the gradient Guadeloupe nel was only created by the reinforced transpar- rainforest: excelsa Vahl, Amanoa ca- Dacryodes ent plastic cover which caused a more signifi- ribaea Kr et Urb and Podocarpus coriaceus LC cant warming effect. Rich are late successional shade-tolerant spe- The was to all the cies; Simaruba amara Aubl and Richeria grandis protocol applied species P coriaceus and A caribaea. The P coria- Vahl are middle successional shade-intolerant except species; Byrsonima coriacea is present in mid- ceus seedlings were placed under the same dle and late succession, whereas Symphonia moderately shaded tunnel (tunnel III) in March 1981 and then to the different globulifera L, a wet soil specialist, is a late suc- subjected experi- mental conditions in 1982. The cessional species. However, their shade reac- January experi- tion is not well known. ment with A caribaea started in March 1982. In each were D excelsa and S amara have compound tunnel, plants grouped by spe- cies with a container of 16 m2. leaves, while the other species have simple density plants per All the were moved a in- leaves. All could be easily identified in the forest plant groups once week side each tunnel so that same understorey with the exception of B coriacea, they occupied the every 8 weeks.
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