On the spatial distribution of woody plant species in a tropical montane cloud forest

Abstract The cloud forest is a special type of forest ecosystem that depends on suitable conditions of humidity and temperature to exist; hence, it is a very fragile ecosystem. The cloud forest is also one of the richest ecosystems in terms of species diversity and rate of endemism. However, today, it is one of the most threatened ecosystems in the world. Little is known about tree species distribution and coexistence among cloud forest trees. Trees are essential to understanding ecosystem functioning and maintenance because they support the ecosystem in important ways. For this dissertation, an analysis of woody plant species distribution at a small scale in a north-Peruvian Andean cloud forest was performed, and some of the factors implicated in the observed patterns were identified. Towards that end, different natural factors acting on species distribution within the forest were investigated: (i) intra-specific arrangements, (ii) heterospecific spatial relationships and (iii) relationships with external environmental factors. These analyses were conducted first on standing woody plants and then on seedlings. The woody plants were found to be clumped in the forest, either considering all the species together or each species separately. However, each species presented a specific pattern and specific spatial relationship among different-age individuals. Dispersal mode, growth form and shade tolerance played roles in the final distribution of the species. Furthermore, spatial associations among species, either positive or negative, were observed. These associations were more numerous when considering individuals of the interacting species at different developmental stages, i.e., younger individuals from one species and older individuals from another. Accordingly, competition and facilitation are asymmetric processes and vary throughout the life of an individual. Moreover, some species appear to prefer certain habitat conditions and avoid other habitats. The habitat definition that best explains species distribution is that which includes both environmental and stand characteristics; thus, a combination of these factors is necessary to understanding species' niche preferences. Seedling distribution was also associated with habitat conditions, but these conditions explained less than the 30% of the spatial variation. The position of conspecific adult individuals also affected seedling distribution; although the seedlings of many tree species avoid the vicinity of conspecifics, a few species appeared to prefer the formation of cohorts around their parent trees. The importance of habitat conditions and distance dependence with conspecifics varied among regions within the forest as well as on the developmental stage of the stand. The results from this thesis suggest that different species can coexist within a given space, forming a “puzzle” of species as a result of the intra- and interspecific spatial relationships along with niche preferences and adaptations that operate at different scales. These factors not only affect each species in a different way, but specific preferences also vary throughout species' lifespans. Resumen Resumen El bosque de niebla es uno de los ecosistemas más amenazados del mundo además de ser uno de los más frágiles. Son formaciones azonales que dependen de la existencia de unas condiciones de humedad y temperatura que permitan la formación de nubes que cubran el bosque; lo que dificulta en gran medida su conservación. También es uno de los ecosistemas con mayor riqueza de especies además de tener uno de los mayores porcentajes de endemismos. Uno de los aspectos más importantes para entender el ecosistema, es identificar y entender los elementos que lo componen y los mecanismos que regulan las relaciones entre ellos. Los árboles son el soporte del ecosistema. Sin embargo, apenas hay información sobre la distribución y coexistencia de los árboles en los bosques de niebla. Esta tesis presenta un análisis de la distribución a pequeña escala de las plantas leñosas en un bosque de niebla situado en la cordillera andina del norte de Perú; así como el análisis de algunos de los factores que pueden estar implicados en que se origine la distribución observada. Para este propósito se estudia cómo influyen factores de diferente naturaleza en la distribución de las especies (i) organización intra-específica (ii) relaciones espaciales heterospecíficas y (iii) relación con factores ambientales externos. En estos análisis se estudiaron primero las plantas jóvenes y las adultas, y después las plántulas. Los árboles aparecieron agregados en el bosque, tanto considerando todos a la vez como cuando se estudió cada especie por separado. Sin embargo, cada especie mostró un patrón distinto así como una particular relación espacial entre individuos jóvenes y adultos. El modo de dispersión, la forma de vida y la tolerancia de la especies estuvieron relacionados con el patrón general observado. Se vio también que ciertas especies aparecían relacionadas con otras, tanto de forma positiva (compartiendo zonas) como negativa (apareciendo en áreas distintas). Las asociaciones fueron mucho más numerosas cuando se consideraron los pares de especies en diferente estado de desarrollo, es decir, individuos jóvenes de una especie e individuos mayores de la otra. Eso indicaría que los procesos de competencia y facilitación son asimétricos y además varían durante la vida de la planta. Por otro lado, algunas especies aparecen preferentemente bajo ciertas condiciones de hábitat y evitan otras. La definición de hábitat a la que mejor responden las especies es cuando se incluyen tanto variables ambientales como de masa; así que ambos tipos de variables son necesarias para entender la preferencia de las especies por ciertos nichos. La distribución de las plántulas también estuvo relacionada con condiciones de hábitat, pero eso sólo llegaba a explicar hasta un 30% de la variabilidad espacial. La posición de los adultos de la misma especie también afectó a la distribución de las plántulas. En bastantes especies las plántulas evitan la cercanía de adultos de su misma especie, padres potenciales, aunque algunas especies aisladas mostraron el patrón contrario y aparecieron preferentemente en las mismas áreas que sus padres. La importancia de las condiciones de hábitat y posición de los adultos en la disposición de las plántulas varía de una zona a otra del bosque y además también varía según el estado de desarrollo de la masa.

Wettability, polarity and water absorption of Quercus ilex leaves: effect of leaf side and age

Plant trichomes play important protective functions and may have a major influence on leaf surface wettability. With the aim of gaining insight into trichome structure, composition and function in relation to water-plant surface interactions, we analyzed the adaxial and abaxial leaf surface of Quercus ilex L. (holm oak) as model. By measuring the leaf water potential 24 h after the deposition of water drops on to abaxial and adaxial surfaces, evidence for water penetration through the upper leaf side was gained in young and mature leaves. The structure and chemical composition of the abaxial (always present) and adaxial (occurring only in young leaves) trichomes were analyzed by various microscopic and analytical procedures. The adaxial surfaces were wettable and had a high degree of water drop adhesion in contrast to the highly unwettable and water repellent abaxial holm oak leaf sides. The surface free energy, polarity and solubility parameter decreased with leaf age, with generally higher values determined for the abaxial sides. All holm oak leaf trichomes were covered with a cuticle. The abaxial trichomes were composed of 8% soluble waxes, 49% cutin, and 43% polysaccharides. For the adaxial side, it is concluded that trichomes and the scars after trichome shedding contribute to water uptake, while the abaxial leaf side is highly hydrophobic due to its high degree of pubescence and different trichome structure, composition and density. Results are interpreted in terms of water-plant surface interactions, plant surface physical-chemistry, and plant ecophysiology.

Plant β-diversity i in human-altered forest ecosystems: the importance of the structural, spatial, and topographical characteristics of stands in patterning plant species assemblages

An understanding of spatial patterns of plant species diversity and the factors that drive those patterns is critical for the development of appropriate biodiversity management in forest ecosystems. We studied the spatial organization of plants species in human- modified and managed oak forests (primarily, Quercus faginea) in the Central Pre- Pyrenees, Spain. To test whether plant community assemblages varied non-randomly across the spatial scales, we used multiplicative diversity partitioning based on a nested hierarchical design of three increasingly coarser spatial scales (transect, stand, region). To quantify the importance of the structural, spatial, and topographical characteristics of stands in patterning plant species assemblages and identify the determinants of plant diversity patterns, we used canonical ordination. We observed a high contribution of ˟-diversity to total -diversity and found ˟-diversity to be higher and ˞-diversity to be lower than expected by random distributions of individuals at different spatial scales. Results, however, partly depended on the weighting of rare and abundant species. Variables expressing the historical management intensities of the stand such as mean stand age, the abundance of the dominant tree species (Q. faginea), age structure of the stand, and stand size were the main factors that explained the compositional variation in plant communities. The results indicate that (1) the structural, spatial, and topographical characteristics of the forest stands have the greatest effect on diversity patterns, (2) forests in landscapes that have different land use histories are environmentally heterogeneous and, therefore, can experience high levels of compositional differentiation, even at local scales (e.g., within the same stand). Maintaining habitat heterogeneity at multiple spatial scales should be considered in the development of management plans for enhancing plant diversity and related functions in human-altered forests