Changes in structure and composition of evergreen forests on an altitudinal gradient in the Venezuelan Guayana Shield
Lionel Hernández, Nelda Dezzeo, Elio Sanoja, Leandro Salazar & Hernán Castellanos
ABSTRACT
There have been several ecological studies in forests of the Guayana Shield, but so far none had examined the changes in structure and composition of evergreen forests with altitude. This study describes and analyzes the structure, species composition and soil characteristics of forest stands at different altitudinal zones in Southeastern Venezuelan Guayana, in order to explain the patterns and the main factors that determine the structure and composition of evergreen forests along the altitudinal gradient. Inventories of 3 948 big (>10cm DBH) and 1 328 small (5-10cm DBH) woody stems were carried out in eleven plots, ranging from 0.1 to 1.0ha, along a 188km long transect with elevations between 290 and 1 395masl. It has been found that 1) hemiepihytes become more dominant and lianas reduce their dominance with increasing altitude and 2) the forest structure in the study area is size-dependent. Five families and 12 genera represented only 9% of the total number of families and genera, respectively, recorded troughout the gradient, but the two groups of taxa comprised more than 50% of the Importance Value (the sum of the relative density and the relative dominance) of all measured stems. Moreover, the results suggest that low species richness seems to be associated with the dominance of one or few species. Stand-level wood density (WD) of trees decreased significantly with increasing elevation. WD is an indicator of trees’life history strategy. Its decline suggests a change in the functional composition of the forest with increasing altitude. The Canonical Correspondence Analysis (CCA) indicated a distinction of the studied forests on the basis of their altitudinal levels and geographic location, and revealed different ecological responses by the forests, to environmental variables along the altitudinal gradient. The variation in species composition, in terms of basal area among stands, was controlled primarily by elevation and secondarily by rainfall and soil conditions. There are other interacting factors not considered in this study like disturbance regime, biological interactions, productivity, and dispersal history, which could affect the structure and composition of the forests in the altitudinal gradient. In conclusion, it appears that the structural and floristic variability observed in the studied transect is produced by a combination of different climates and randomly expressed local processes interacting across a complex physical landscape.
RESUMEN
A pesar de los diversos estudios ecológicos realizados en los bosques del Escudo de Guayana, ninguno de ellos había analizado hasta ahora los cambios en la composición y estructura de bosques siempreverdes que ocurren al incrementar la altitud. Con el fin de identificar patrones y factores determinantes de la estructura y la composición de bosques en un gradiente altitudinal (290-1 395msnm) en el sudeste de Venezuela se realizaron inventarios de 3 948 fustes grandes (>10cm DAP) y 1 328 pequeños (5-10cm DAP) en once parcelas a lo largo de un transecto de 188km. Con el incremento de la altitud, el área basal aumenta en las hemiepífitas y disminuye en las lianas. Las familias y los géneros más importantes representaron una reducida proporción del total de taxones, pero abarcaron más del 50% del Valor de Importancia. Bajos valores de riqueza se asocian con la dominancia de pocas especies. La densidad de madera decrece significativamente a mayor altitud, y sugiere un cambio en la composición funcional. Existen diferentes respuestas ecológicas de los rodales ante cambios físico-ambientales. La variación de la composición de especies en el gradiente fue controlada por altitud, lluvia y condiciones edáficas; pero se desconoce el efecto de otros factores no considerados como régimen de perturbaciones, interacciones biológicas, productividad e historia de dispersión. La variabilidad observada en la estructura y la composición de bosques en el gradiente parece ser un resultado del efecto combinado de diferentes climas y procesos locales aleatorios que interactúan en un complejo paisaje.