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On the Spatial Distribution of Woody Plant Species in a Tropical Montane Cloud Forest

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On the Spatial Distribution of Woody Plant Species in a Tropical Montane Cloud Forest UNIVERSIDAD POLITÉCNICA DE MADRID ESCUELA TÉCNICA SUPERIOR DE INGENIEROS DE MONTES ON THE SPATIAL DISTRIBUTION OF WOODY PLANT SPECIES IN A TROPICAL MONTANE CLOUD FOREST TESIS DOCTORAL ALICIA LEDO ÁLVAREZ Ingeniera de Montes MADRID, 2012 PROGRAMA DE DOCTORADO DE ECONOMÍA Y GESTIÓN FORESTAL ESCUELA TÉCNICA SUPERIOR DE INGENIEROS DE MONTES UNIVERSIDAD POLITÉCNICA DE MADRID ON THE SPATIAL DISTRIBUTION OF WOODY PLANT SPECIES IN A TROPICAL MONTANE CLOUD FOREST ALICIA LEDO ÁLVAREZ Ingeniera de Montes DIRECTORES SONIA CONDÉS RUIZ FERNANDO MONTES PITA Doctora Ingeniera de Montes Doctor Ingeniero de Montes MADRID, 2012 Tribunal nombrado por el Mgfco. y Excmo. Sr. Rector de la Universidad Politécnia de Madrid, el día…………de……………………..de 2012 Presidente D. …………………………………………………….. Vocal D. …………………………………………………….. Vocal D. …………………………………………………….. Vocal D. …………………………………………………….. Secretario D. …………………………………………………….. Realizando el acto de defensa y lectura de la Tesis el día ……….de…….…….. de 2012 en Madrid Calificación………………….. EL PRESIDENTE LOS VOCALES EL SECRETARIO Abstract 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. Contents Contents i Thesis Contents Chapter I Introduction Chapter II Research Goals Chapter III Cloud Forest Review Chapter IV Overview of the Employed Methods Chapter V Study Site and Inventory Chapter VI Dasometric and Environmental Values of the Experimental Plots Chapter VII The Spatial Organisation of the Woody Plant Species Chapter VIII The Spatial Relationships among Woody Plant Species Chapter IX The Influence of Habitat Conditions on Woody Plant Species Distribution Chapter X Recruitment Patterns Chapter XI Synthesis and Discussion Chapter XII Conclusions ii Resumen de la tesis en castellano iii Appendix CHAPTER I Introduction Chapter I: Introduction 1. Introduction The cloud forest is one of the most threatened ecosystems in the world (Hamilton 1995; Foster 2001; Bruijnzeel et al. 2011). It is also one of the most fragile (Gomez-Peralta et al. 2008), making its conservation difficult. Cloud forests are threatened not only due to direct human influence, such as logging and soil conversion (Sarmiento 1993; Bubb et al. 2004; Aubad et al. 2008), but also by indirect effects, such as changes detected in the microclimatic conditions necessary for cloud forest maintenance (Ledo et al. 2009), which is likely a direct consequence of climate change (Pounds et al. 1999; Still et al. 1999; Foster 2001). The cloud forest is an azonal ecosystem, and it depends sine qua non on the convergence of specific environmental characteristics (Foster 2001). Therefore, due to their specific distribution, cloud forests have been identified as “islands” of vegetation throughout the world (Howard 1970). The most important requirements for cloud forest formation are high humidity and a suitable temperature that results in clouds or fog being present in the forest for most of the day over most or the whole of the year. Fog formation depends mainly on geomorphologic conditions and less on climatic conditions. As a result, cloud forests are found mainly on mountain slopes in the Central Mountains
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