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Stand Structure of Monocotyledons and Dicotyledons in Different BOIS ET FORÊTS DES TROPIQUES, 2011, N° 307 (1) STRUCTURE DE PEUPLEMENTS / LE POINT SUR… 33 Stand structure of Monocotyledons and Dicotyledons in different successional stages in Corcovado Fidèle Bognounou1, 2, 3 Renée Morton3 National Park, Costa Rica Sarah Ayangma3 Laurence Jonkers3 Christer Björkman3 Helena Bylund3 Colin M. Orians4 Andres Vega5 Per Chister Oden2 1 University of Ouagadougou Life Sciences and Earth Sciences Training and Research Department 03 BP 7021, Ouagadougou 03 Burkina Faso 2 Swedish University of Agricultural Sciences Faculty of Forest Sciences Southern Swedish Forest Research Centre PO Box 101, SE-230 53 Alnarp Sweden 3 Swedish University of Agricultural Sciences Faculty of Forest Sciences Department of Ecology Box 7044, 75007 Uppsala Sweden 4 Department of Biology Tufts University 120 Dana Building Medford, MA 02155 USA 5 Corcovado National Park 400 E, 75 S, 75 E de la Municipalidad de Tibas Tibas Costa Rica Photograph 1. Primary forest. Photograph S. Ayangma. BOIS ET FORÊTS DES TROPIQUES, 2011, N° 307 (1) F. Bognounou, R. Morton, 34 FOCUS / STAND STRUCTURE S. Ayangma, L. Jonkers, C. Björkman, H. Bylund, C. M. Orians, A. Vega, P. C. Oden RÉSUMÉ ABSTRACT RESUMEN STRUCTURE DES PEUPLEMENTS DE STAND STRUCTURE OF MONOCOTYLEDONS ESTRUCTURA DE LOS RODALES DE MONOCOTYLEDONES ET DICOTYLEDONES AND DICOTYLEDONS IN DIFFERENT MONOCOTILEDÓNEAS Y DICOTILEDÓNEAS LORS DE DIFFERENTS STADES DE SUCCESSIONAL STAGES IN CORCOVADO EN DIFERENTES ETAPAS DE SUCESIÓN SUCCESSION DANS LE PARC NATIONAL NATIONAL PARK, COSTA RICA EN EL PARQUE NACIONAL DE CORCOVADO DE CORCOVADO AU COSTA RICA EN COSTA RICA Les deux grands groupes d’angiospermes, The two major groups of angiosperms, Los dos principales grupos de angiosper- monocotylédones et dicotylédones, diffè- monocotyledons and dicotyledons, differ mas, monocotiledóneas y dicotiledóneas, rent à plusieurs égards au niveau de leur in several of their life history traits. difieren en varios aspectos de su ciclo cycle de vie. Malgré les différences évi- Despite the obvious differences between vital. A pesar de las evidentes diferencias dentes entre ces deux groupes, peu the two groups, there are few studies entre estos dos grupos, existen pocos d’études se sont penchées de façon appro- elaborating on how these may relate to estudios que hayan profundizado en fondie sur la manière dont cela pourrait their ecological succession. The study cómo podría esto relacionarse con su influencer leur succession écologique. compares the relative densities of these sucesión ecológica. Éste estudio compara L’étude compare les densités relatives de two fundamental groups of plants with las densidades relativas de estos dos gru- ces deux groupes fondamentaux à l’évolu- different evolutionary histories in forests pos fundamentales de plantas de distinta tion contrastée, lors de deux stades de suc- in two different successional stages at evolución, en dos etapas de sucesión cession forestière à la station de Sirena the Sirena station in Costa Rica’s Corcov- forestal en la estación de Sirena en el Par- dans le Parc national de Corcovado au ado National Park, in Central America. que Nacional de Corcovado, en Costa Costa Rica, en Amérique centrale. Selon The hypothesis was that the monoco - Rica. Según ésta hipótesis, la relación l'hypothèse de départ, le rapport monoco- tyledons/dicotyledons ratio would monocotiledóneas/dicotiledóneas debe- tylédones/dicotylédones devrait diminuer decrease in later successional stages. ría disminuir en las etapas más avanzadas aux stades plus avancés de la succession. A total of 40 right-angled triangular plots de la sucesión. Se muestrearon un total L’échantillonnage a porté sur un ensemble ((5 ×5m)/2) were sampled and the de 40 parcelas triangulares de ángulo de 40 placettes triangulaires à angle droit plants were divided into four crown size recto ((5 × 5 m)/2), en las que se clasifica- ((5 × 5 m)/2), où les plantes ont été clas- classes, as well as into monocotyledons ron las plantas en cuatro clases según el sées en quatre catégories selon la dimen- and dicotyledons. Plant densities and tamaño de las copas, así como en mono- sion des houppiers, ainsi qu’en monocoty- crown size class distribution were cotiledóneas y dicotiledóneas. Se efectuó lédones et dicotylédones. Une analyse de analysed using ANOVA with the general un análisis de la distribución de densida- la distribution des densités et des classes linear model (GLM) procedure. The mean des y clases de tamaño de las copas de dimension des houppiers a été effectuée overall plant density in the secondary for- mediante ANOVA (modelo lineal general). par ANOVA selon la procédure de modélisa- est was significantly greater than in the La densidad promedio total del bosque tion linéaire généralisée. La densité primary forest (P < 0.001). The structure secundario fue significativamente mayor moyenne totale de la forêt secondaire of the crown size class distribution, how- que la del bosque primario (P < 0.001). Sin apparaît significativement plus importante ever, was similar for both monocotyle- embargo, la estructura de distribución de que celle de la forêt primaire (P < 0,001). dons and dicotyledons, and for the pri- las clases de tamaño de las copas fue Cependant, la structure de la distribution mary and secondary forest types. There similar entre monocotiledóneas y dicotile- des classes de dimension des houppiers was no significant difference between dóneas, y también entre bosque primario est proche entre monocotylédones et dico- the two forests in the monocotyledons to y secundario. No aparece ninguna diferen- tylédones, mais aussi entre forêt primaire et dicotyledons ratio (P = 0.99). The crown cia significativa entre los dos tipos de secondaire. Il n’apparaît aucune différence size class distribution results indicate bosque en la relación monocotiledó- significative entre les deux types de forêt that there is a similar distribution trend neas/dicotiledóneas (P = 0.99). Los resul- dans le rapport monocotylédones / dicoty- in monocotyledon and dicotyledon suc- tados de la distribución de clases de lédones (P = 0,99). Les résultats concernant cession, although there are always more tamaño de las copas indican una tenden- la distribution des classes de dimension dicotyledons than monocotyledons. cia similar en la distribución de sucesio- des houppiers indiquent une tendance These results led to conclude that a nes de monocotiledóneas y dicotiledó- similaire dans la distribution des succes- larger difference between successional neas, aunque las dicotiledóneas sean sions de monocotylédones et dicotylé- stages is necessary to perceive the differ- siempre más numerosas. Estos resultados dones, même si les dicotylédones sont tou- ence anticipated in this study. llevan a la conclusión de que las diferen- jours plus nombreux. Ces résultats amè- cias entre etapas de sucesión deben ser nent à conclure qu'il est nécessaire d'avoir Keywords: rain forest, primary forest, sec- mayores para percibir la diferencia antici- une importante différence entre les stades ondary forest, plant evolution, humid trop- pada en éste estudio. de succession avant de percevoir la diffé- ics, Central America. rence anticipée dans cette étude. Palabras clave: bosque tropical húmedo, bosque primario, bosque secundario, evo- Mots clé : forêt tropicale humide, forêt pri- lución florística, trópicos húmedos, América maire, forêt secondaire, évolution floristique, Central. tropiques humides, Amérique centrale. BOIS ET FORÊTS DES TROPIQUES, 2011, N° 307 (1) STRUCTURE DE PEUPLEMENTS / LE POINT SUR… 35 INTRODUCTION Flowering plants are conventionally, divided into two major classes: Dicotyledons (Magnoliopsida) and Monocotyledons (Liliopsida). This separation into two classes is commonly taken for granted, because it is “patently obvious”, but botanists have not always recog- nized these as the two fundamental groups of angiosperms (Barabé, Brouillet, 1982). Despite the problems with rec- ognizing basal angiosperm taxa, the conventional distinc- tions between dicotyledons and monocotyledons are still quite useful. The main morphological differences between monocotyledons and dicotyledons are, respectively, embryo with a single cotyledon vs. embryo with two cotyle- dons; pollen with a single furrow or pore vs. pollen with three furrows or pores; flower parts in multiples of three vs. flower parts in multiples of four or five; parallel major leaf veins vs. reticulated major leaf vein; scattered vs. ring- shaped pattern of stem vascular bundles; adventitious Photograph 2. roots vs. roots developing from radicles; secondary growth Secondary forest. absent vs. secondary growth often present. It is thanks to Photograph F. Bognounou. the extreme plasticity of their vegetative and reproductive organs that angiosperms have become so widely and suc- cessfully established; their genesis was the ultimate link in the largest remaining tract of rainforest on the Pacific coast the ongoing chain of evolution of our planet’s plant of Central America (Janzen et al., 1985; Vaughan, 1981). So kingdom (Takhtajan, 1953). It is universally agreed that far, over 1,510 vascular plant species in 707 genera and 154 monocotyledons are derived from primitive dicotyledons, families have been documented (Fnt, 1992). Based on suc- and that monocotyledons must follow rather than precede cessional stages, the park was divided into two different for- dicotyledons in any proper linear sequence (Cronquist, est types, primary and secondary forests. The primary forest 1965). Few conclusions
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