EMBRAPA Amazônia Ocidental, Manaus, Brazil. Universität Hamburg, Deutschland

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1 Morphological Traits and Ecological Behaviour of Selected Secondary Forest Tree Species in the Central Amazon Aguiar, M. O. 1, de Morais, R. R. 1, Araújo, R. da C. 1, Skatulla, M. 2 and Preisinger, H. 2 1 EMBRAPA Amazônia Ocidental, Manaus, Brazil 2 Universität Hamburg, Deutschland 1 Introduction Prognoses on vegetation development after severe disturbance events like slashing and burning require a good knowledge of the autecological behavior of the plant species involved in the successional processes. This is not yet available for Amazonian plant species. Thirteen common secondary forest tree species of Terra Firme sites near Manaus, Amazonas, Brazil, covering a wide range of growth-form types and of ecological behavior in the succcessional sequence, were selected for a comparative study designed to develop basic knowledge of their autecology. Growth-form, biometric traits and the morphological and anatomical characteristics of leaves and wood were compared. Plant biomass and the mineral nutrient content of different organs were analyzed in individual plants. In addition, the life history of individual twigs and leaves was recorded. In the approach we proceed from the well-known fact that growth-forms of vascular plants represent a complex of characteristics closely linked to the ecological behavior of the species and their site conditions. The aim of the study is to identify functional traits for plant species in the secondary forest and to derive "plant functional types" for different successional stages. The present work searches for causal links between growth-form, leaf characteristics and behavior of thirteen sample species of Central Amazonian secondary forests. reference, with regard to the growth-form spectrum and the frequency of the species, is displayed in Fig Morphological traits of leaves The majority of the plant species occurring in the secondary forest shows a large uniformity in leaf shape, having simple, ovate or lanceolate leaves (cf. Fig. 1). However, in detail 2 Sample species The sample species (see Fig. 1) and the individuals for study were selected from a 1 ha secondary forest site which served as an area for reference. Important criteria for the choices were: 1 availability of a large number of plant individuals of the species, 2 availability of some ecological information on the species, either from literature or from own observations, and 3 the species represent examples for a certain ecological type in the secondary forest. The role of the sample species in the secondary forest for Fig. 1: Sample species, growth-form types and leaves there is a high diversity in leaf shape, concerning the leaf apex (e.g. obtuse, acuminate, aristate) and the leaf margins (e.g. entire, dentate, serrate). Some species have lobed, palmate or digitate leaves (e.g. Pouroma spp., Cecropia spp., Schefflera spp.). Even more striking than the diversity in leaf shape are the differences in leaf size (leaf areas of the sample species between approx.10 cm 2 and cm 2, cf. Fig. 1) and the diversity in the structure of leaf surfaces (glabrous, papillose, ± hairy, felty, cf. Tab. 1 and Fig. 3). 965

2 German-Brazilian Workshop on Neotropical Ecosystems Achievements and Prospects of Cooperative Research Fig. 2: Sample species and their occurrence in the spectra of species frequencies (left) and growth-form types (right) of the secondary forest under study. WT = Lianas, TT = Tall Trees, LT = Low Trees, MT = Medium sized Trees, ST = Treelets. 4 Morphological traits, biomass and ecological behavior In the analysis, three plant individuals of each of thirteen sample species were examined with regard to overground plant biomass (dry matter), leaf biomass per plant, mean values of leaf areas (from 50 leaves each) and specific leaf area weight. The resulting two ordination diagrams of the individuals of the sample species reveal the following trends (Figs. 4 and 5): The proportion of leaf biomass on overground biomass decreases with growing overground plant biomass (see Fig. 4). There are large-leaved species with high and with low specific leaf area weights (Bellucia spp. vs. Miconia tomentosa and Cecropia concolor) in the analysis. The small-leaved species Bocageopsis multiflora and Goupia glabra have low leaf area weights. Miconia alata, a plant species with medium-sized leaves, has a low leaf area weight. All the other species occupy intermediate positions in the diagram. The large leaves are hairy, and the small leaves tend to be glabrous. The combinations of traits mentioned are linked to certain growth-form types. The progressive succession of secondary forest tree species commences with the large-leaved, short lived sample species (Treelets) and progresses to small-leaved, longer-lived species (Medium and Tall trees). The many liana species Tab. 1: Sample species and characteristics of leaves 966

3 glabrous (Bocageopsis multiflora). These sets of traits indicate different strategies for an efficient use of resources in a changing environment during a progressive succession. Type 1 is fast growing, short lived (± 10 years), invests little energy in trunk and twigs and appears early in succession. The hairy leaf surface might protect the leaves from drought in the open landscape. Type 3 invests much energy in a durable trunk and twigs, but little in the single leaf. The many, small leaves reduce the risk of loosing a considerable part of the canopy because of the destruction of single leaves. Type 2 are low and medium sized trees which invest much energy in large and durable leaves. The leaves are well protected against phytophages and mechanical damage. The majority of studies on "plant functional types and disturbance dynamics" refers to a large geographic scale or the global scale (cf. MCINTYRE et al. 1999). The present study identified functional traits and types on a local scale. The contrasting types identified in the example occur in the same sites, either at the same time or at different times in the successional sequence. This study is based on results of the SHIFT-Project "Recultivation of degraded areas" (ENV 23), subproject "Vegetation", carried out by EMBRAPA Amazônia Ocidental, Manaus and the Institute of Applied Botany, University of Hamburg. 6 References Fig. 3: Sample species and leaf surfaces show a high plasticity with regard to the analyzed traits and are therefore abundant in a large spectrum of sites. MCINTYRE, S., DÍAZ, S., LAVOREL, S. & CRAMER, W Plant functional types and disturbance dynamics - Introduction. J. Vegetation Science 10/5, PREISINGER, H., DÜNISCH, O., MORAIS, R.R. DE, RICHTER, K. & ARAÚJO, R. DA C Comparitive studies on morpho-physiological traits of six Amazonian species of Bellucia and Miconia (Melastomataceae) and implications for their ecological behaviour. Mitt. BFH Hamburg 193, Conclusions The analysis reveals at least three fundamentally different ecological types of secondary forest tree species and some of their functional traits (example species in brackets): Type 1: Treelet, fraction leaf biomass / overground biomass high - ± large leaves - low leaf area weight - leaves hairy (Miconia tomentosa); Type 2: Low tree, large leaves - high leaf area weight - leaves ± hairy (Bellucia dichotoma); Type 3: Medium tree, fraction leaf biomass / overground biomass low - small leaves - low leaf area weight - leaves 967

4 Fig. 4: Ordination of sample species and individuals with regard to overground plant biomass and leaf biomass as proportion of overground biomass Fig. 5: Ordination of sample species with regard to specific leaf area weight (weight per leaf area = reciprocal value of leaf area index) and leaf area 968

5 Editors Reinhard Lieberei 1, Helmut K. Bianchi 2, Vera Boehm 1, Christoph Reisdorff 1 1 Universität Hamburg, Institut für Angewandte Botanik, Ohnhorststr. 18, Hamburg, Germany 2 GKSS-Forschungszentrum Geesthacht GmbH, Max-Planck-Straße 1, Geesthacht Germany Layout Printing Helmut K. Bianchi, GKSS, Karsten Bittner, Documedia, Geesthacht, Germany GKSS-Forschungszentrum Geesthacht GmbH, Geesthacht, Germany ISBN X Lieberei, R., Bianchi, H-K., Boehm, V., Reisdorff, C., (eds.) 2002: Neotropical Ecosystems, Proceedings of the German-Brazilian Workshop, Hamburg GKSS-Geesthacht. The publishers give permission to copy single contributions from the Proceedings for personal use exclusively. Copies may be passed on only with the correct specification of the source. The research cooperation has been carried out under the auspices of the German - Brazilian Govemmental Agreement on the Cooperation in Scientific Research and Technological Development. The issuance of the Proceedings and the production of the CD-ROM was sponsored (Code ) by the Federal Ministry of Education and Research The responsibility for the contents of the contributions is solely the authors'.

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