Deforestación y Pérdida de hábitat en Bosques de montaña en la Cuenca alta del Río Zamora (Loja, Ecuador)

Los bosques húmedos de montaña se encuentran reconocidos como uno de los ecosistemas más amenazados en el mundo, llegando inclusive a ser considerado como un “hotspot” por su alta diversidad y endemismo. La acelerada pérdida de cobertura vegetal de estos bosques ha ocasionado que, en la actualidad, se encuentren restringidos a una pequeña fracción de su área de distribución histórica. Pese a esto, los estudios realizados sobre cual es efecto de la deforestación, fragmentación, cambios de uso de suelo y su efecto en las comunidades de plantas presentes en este tipo de vegetación aún son muy escuetos, en comparación a los realizados con sus similares amazónicos. En este trabajo, el cual se encuentra dividido en seis capítulos, abordaremos los siguientes objetivos: a) Comprender cuál es la dinámica que han seguido los diferentes tipos de bosques montanos andinos de la cuenca del Rio Zamora, Sur de Ecuador durante entre 1976 y 2002. b) Proveer de evidencia de las tasas de deforestación y fragmentación de todos los tipos diferentes de bosques montanos andinos presentes en la cuenca del Rio Zamora, Sur de Ecuador entre 1976 y 2002. c) Determinar qué factores inducen a la fragmentación de bosques de montaña en la cuenca alta del río Zamora entre 1976 y 2002. d) Determinar cuáles son y cómo afectan los factores ambientales y socioeconómicos a la dinámica de la deforestación y regeneración (pérdida y recuperación del hábitat) sufrida por los bosques de montaña dentro de la zona de estudio y e) Determinar si la deforestación y fragmentación actúan sobre la diversidad y estructura de las comunidades de tres tipos de organismos (comunidades de árboles, comunidades de líquenes epífitos y comunidades de hepáticas epífitas). Este estudio se centró en el cuenca alta del río Zamora, localizada al sur de Ecuador entre las coordenadas 3º 00´ 53” a 4º 20´ 24.65” de latitud sur y 79º 49´58” a 78º 35´ 38” de longitud oeste, que cubre alrededor de 4300 km2 de territorio situado entre las capitales de las provincias de Loja y Zamora-Chinchipe. Con objeto de predecir la dinámica futura de la deforestación en la región de Loja y cómo se verán afectados los diferentes tipos de hábitat, así como para detectar los factores que más influyen en dicha dinámica, se han construido modelos basados en la historia de la deforestación derivados de fotografías aéreas e imágenes satelitales de tres fechas (1976, 1989 y 2002). La cuantificación de la deforestación se realizó mediante la tasa de interés compuesto y para la caracterización de la configuración espacial de los fragmentos de bosque nativo se calcularon índices de paisaje los cuales fueron calculados utilizando el programa Fragstats 3.3. Se ha clasificado el recubrimiento del terreno en forestal y no forestal y se ha modelado su evolución temporal con Modelos Lineales Generalizados Mixtos (GLMM), empleando como variables explicativas tanto variables ambientales espacialmente explícitas (altitud, orientación, pendiente, etc) como antrópicas (distancia a zonas urbanizadas, deforestadas, caminos, entre otras). Para medir el efecto de la deforestación sobre las comunidades modelo (de árboles, líquenes y hepáticas) se monitorearon 11 fragmentos de vegetación de distinto tamaño: dos fragmentos de más de cien hectáreas, tres fragmentos de entre diez y noventa ha y seis fragmentos de menos de diez hectáreas. En ellos se instalaron un total de 38 transectos y 113 cuadrantes de 20 x 20 m a distancias que se alejaban progresivamente del borde en 10, 40 y 80 m. Nuestros resultados muestran una tasa media anual de deforestación del 1,16% para todo el período de estudio, que el tipo de vegetación que más alta tasa de destrucción ha sufrido, es el páramo herbáceo, con un 2,45% anual. El análisis de los patrones de fragmentación determinó un aumento en 2002 de más del doble de fragmentos presentes en 1976, lo cual se repite en el análisis del índice de densidad promedio. El índice de proximidad media entre fragmentos muestra una reducción progresiva de la continuidad de las áreas forestadas. Si bien las formas de los fragmentos se han mantenido bastante similares a lo largo del período de estudio, la conectividad entre estos ha disminuido en un 84%. Por otro lado, de nuestros análisis se desprende que las zonas con mayor probabilidad de deforestarse son aquellas que están cercanas a zonas previamente deforestadas; la cercanía a las vías también influye significativamente en la deforestación, causando un efecto directo en la composición y estructura de las comunidades estudiadas, que en el caso de los árboles viene mediado por el tamaño del fragmento y en el caso del componente epífito (hepáticas y líquenes), viene mediado tanto por el tamaño del fragmento como por la distancia al borde del mismo. Se concluye la posibilidad de que, de mantenerse esta tendencia, este tipo de bosques desaparecerá en corto tiempo y los servicios ecosistémicos que prestan, se verán seriamente comprometidos. ABSTRACT Mountain rainforests are recognized as one of the most threatened ecosystems in the world, and have even come to be considered as a “hotspot” due to their high degree of diversity and endemism. The accelerated loss of plant cover of these forests has caused them to be restricted today to a small fraction of their area of historic distribution. In spite of this, studies done on the effect of deforestation, fragmentation, changes in soil use and their effect on the plant communities present in this type of vegetation are very brief compared to those done on their analogues in the Amazon region. In this study, which is divided into six chapters, we will address the following objectives: a) To understand what the dynamic followed by the different types of Andean mountain forests in the Zamora River watershed of southern Ecuador has been between 1976 and 2002. b) To provide evidence of the rates of deforestation and fragmentation of all the different types of Andean mountain forests existing in the upper watershed of the Zamora River between 1976 and 2002. c) To determine the factors that induces fragmentation of all different types of Andean mountain forests existing in the upper watershed of the Zamora River between 1976 and 2002. d) To determine what the environmental and anthropogenic factors are driving the dynamic of deforestation and regeneration (loss and recuperation of the habitat) suffered by the mountain forests in the area of the study and e) To determine if the deforestation and fragmentation act upon the diversity and structure of three model communities: trees, epiphytic lichens and epiphytic liverworts. This study is centered on the upper Zamora River watershed, located in southern Ecuador between 3º 00´ 53” and 4º 20´ 24.65 south latitude and 79º 49´ 58” to 78º 35´ 38” west longitude, and covers around 4,300 km2 of territory located between Loja and Zamora-Chinchipe provinces. For the purpose of predicting the future dynamic of deforestation in the Loja region and how different types of habitats will be affected, as well as detecting the environmental and socioeconomic factors that influence landscape dynamics, models were constructed based on deforestation history, derived from aerial photographs and satellite images for three dates (1976, 1989 and 2002). Quantifying the deforestation was done using the compound interest rate; to characterize the spatial configuration of fragments of native forest, landscape indices were calculated with Fragstats 3.3 program. Land cover was classified as forested and not forested and its evolution over time was modeled with Generalized Linear Mixed Models (GLMM), using spatially explicit environmental variables (altitude, orientation, slope, etc.) as well as anthropic variables (distance to urbanized, deforested areas and roads, among others) as explanatory variables. To measure the effects of fragmentation on three types of model communities (forest trees and epiphytic lichen and liverworts), 11 vegetation fragments of different sizes were monitored: two fragments of more than one hundred hectares, three fragments of between ten and ninety ha and six fragments of fewer than ten hectares . In these fragments, a total of 38 transects and 113 20 x 20 m quadrats were installed at distances that progressively moved away from the edge of the fragment by 10, 40 and 80 m. Our results show an average annual rate of deforestation of 1.16% for the entire period of the study, and that the type of vegetation that suffered the highest rate of destruction was grassy paramo, with an annual rate of 2.45%. The analysis of fragmentation patterns determined the number of fragments in 2002 more than doubled the number of fragments present in 1976, and the same occurred for the average density index. The variation of the average proximity index among fragments showed a progressive reduction of the continuity of forested areas. Although fragment shapes have remained quite similar over the period of the study, connectivity among them has diminished by 84%. On the other hand, it emerged from our analysis that the areas of greatest probability of deforestation were those that are close to previously deforested areas; proximity to roads also significantly favored the deforestation causing a direct effect on the composition of our model communities, that in the case of forest trees is determined by the size of the fragment, and in the case of the epiphyte communities (liverworts and lichens), is determined, by the size of the fragment as well as the distance to edge. A subject under discussion is the possibility that if this tendency continues, this type of forest will disappear in a short time, and the ecological services it provides, will be seriously endangered.

Using ecological niche models to support tree species selection for forest restoration planning in largely deforested regions

Species selection for forest restoration is often supported by expert knowledge on local distribution patterns of native tree species. This approach is not applicable to largely deforested regions unless enough data on pre-human tree species distribution is available. In such regions, ecological niche models may provide essential information to support species selection in the framework of forest restoration planning. In this study we used ecological niche models to predict habitat suitability for native tree species in "Tierra de Campos" region, an almost totally deforested area of the Duero Basin (Spain). Previously available models provide habitat suitability predictions for dominant native tree species, but including non-dominant tree species in the forest restoration planning may be desirable to promote biodiversity, specially in largely deforested areas were near seed sources are not expected. We used the Forest Map of Spain as species occurrence data source to maximize the number of modeled tree species. Penalized logistic regression was used to train models using climate and lithological predictors. Using model predictions a set of tools were developed to support species selection in forest restoration planning. Model predictions were used to build ordered lists of suitable species for each cell of the study area. The suitable species lists were summarized drawing maps that showed the two most suitable species for each cell. Additionally, potential distribution maps of the suitable species for the study area were drawn. For a scenario with two dominant species, the models predicted a mixed forest (Quercus ilex and a coniferous tree species) for almost one half of the study area. According to the models, 22 non-dominant native tree species are suitable for the study area, with up to six suitable species per cell. The model predictions pointed to Crataegus monogyna, Juniperus communis, J.oxycedrus and J.phoenicea as the most suitable non-dominant native tree species in the study area. Our results encourage further use of ecological niche models for forest restoration planning in largely deforested regions.

Forest connectivity restoration through reforestation: an integrated methodology for prioritizing agricultural lands and selecting reforestation species

Forest connectivity restoration is a major goal in natural resource planning. Given the high amount of abandoned cultivated lands, setting efficient methods for the reforestation of agricultural lands offers a good opportunity to face this issue. However, reforestations must be carefully planned, which poses two main challenges. In first place, to determine those agricultural lands that, once reforested, would meet more effectively the planning goals. As a further step, in order to grant the success of the activity, it is fairly advisable to select those tree species that are more adapted to each particular environment. Here we intend to give response to both requirements by proposing a sequential and integrated methodology that has been implemented in two Spanish forest districts, which are formed by several landscape types that were previously defined and characterized. Using the software Conefor Sensinode, a powerful tool for quantifying habitat availability that is based on graph theory concepts, we determined the landscapes where forest planning should have connectivity as a major concern and, afterwards, we detected the agricultural patches that would contribute most to enhance connectivity if they were reforested. The subsequent reforestation species assessment was performed within these priority patches. Using penalized logistic regressions we fitted ecological niche models for the Spanish native tree species. The models were trained with species distribution data from the Spanish Forest Map and used climatic and lithological variables as predictors. Model predictions were used to build ordered lists of suitable species for each priority patch. The lists include dominant and non dominant tree species and allow adding biodiversity goals to the reforestation planning. The result of this combined methodology is a map of agricultural patches that would contribute most to uphold forest connectivity if they were reforested and a list of suitable tree species for each patch ordered by occurrence probability. Therefore the proposed methodology may be useful for suitable and efficient forest planning and landscape designing.

Ecología de la dispersión de plantas en los bosques secos del suroccidente Ecuatoriano

La importancia del proceso de dispersión de semillas en la estructura y dinámica de los ecosistemas es ampliamente reconocida. Sin embargo, para los bosques tropicales estacionalmente secos los estudios relacionados con este proceso son aún escasos y dispersos en comparación con los bosques tropicales lluviosos. En este trabajo se estudió la importancia de los síndromes de dispersión de semillas en la estructuración de comunidades, mediante el análisis de los patrones de dispersión de semillas en el espacio y tiempo para comunidades de leñosas en los bosques secos del suroccidente Ecuatoriano. Esta área forma parte de la región Tumbesina, una de las áreas de endemismo más importantes del mundo, pero también uno de los hotspots más amenazados. El clima se caracteriza por una estación seca que va de mayo a noviembre y una estación lluviosa que se extiende desde diciembre a abril. Para toda esta zona se estima una temperatura promedio anual entre 20° y 26°C y una precipitación promedio anual entre 300 y 700 mm. El trabajo de campo se desarrolló entre febrero de 2009 y septiembre de 2012. El primer paso fue la recopilación de información sobre las especies leñosas nativas de los bosques secos del suroccidente de Ecuador, que permitiera asignar a cada especie a un síndrome de dispersión para determinar el espectro de síndromes de dispersión de semillas. Luego, utilizando la información disponible de 109 parcelas establecidas previamente a lo largo de cuatro cantones de la provincia de Loja que conservan bosques secos en buen estado, se analizó la relación entre el síndrome de dispersión y condiciones ambientales. La relación de los síndromes de dispersión con los patrones espaciales de las especies y con los patrones de la lluvia y banco de semillas se estudió dentro de una parcela permanente de 9 ha, en la Reserva Ecológica Arenillas. Dentro de esta parcela se estableció un transecto de aproximadamente 3,4 km, que se recorrió mensualmente para colectar excretas de cérvidos y analizar el rol de este grupo como dispersor de semillas. Una gran variedad de plantas en los bosques secos tropicales del suroccidente de Ecuador requirió la asistencia de animales para la dispersión de semillas. Sin embargo, un análisis del espectro de dispersión considerando no solo la riqueza, sino también la abundancia relativa de especies, permitió determinar que a pesar de la alta variedad de especies zoócoras, la mayor parte de la comunidad correspondía a individuos anemócoros, que no proveen ninguna recompensa para la dispersión por animales. Este patrón puede deberse a la abundancia relativa de hábitats adecuados para especies con diferente síndrome de dispersión. Las condiciones ambientales afectaron la estructura del espectro de dispersión en la comunidad de bosque seco neotropical estudiada. El análisis de la importancia relativa del síndrome de dispersión y de la heterogeneidad espacial en la formación de patrones espaciales de árboles adultos permitió determinar que la heterogeneidad ambiental ejercía un efecto adicional (y en algunos el único) en la formación de patrones agregados de la mayoría de especies estudiadas. Los resultados señalaron diferencias en los patrones espaciales de las especies dependiendo del síndrome de dispersión, pero también una gran variación en los patrones espaciales incluso entre especies del mismo síndrome de dispersión. El análisis simultáneo de los patrones de la lluvia de semillas y banco de semillas de una comunidad de leñosas y su relación con la vegetación establecida indicaron que la lluvia de semillas era temporalmente variable en número de especies y abundancia de semillas, y dependía del síndrome de dispersión. El síndrome de dispersión también influyó en la formación de bancos de semillas, siendo las especies con capacidad de dispersión limitada (autócoras) las de mayor riqueza de especies y abundancia de semillas. Los cérvidos también se consideraron como un elemento clave en el proceso de dispersión de semillas. Al menos ocho especies leñosas fueron dispersadas legítimamente vía endozoócora. La mayoría de las especies dispersadas presentaron diásporas sin adaptaciones obvias para la dispersión, por lo que la ingestión de semillas por cérvidos se constituye en una vía potencial para la dispersión de sus semillas a largas distancias y, con ello, mejora la posibilidad de colonizar nuevos sitios y mantener el flujo genético. Los resultados de este estudio aportan nuevas evidencias para el entendimiento de la importancia de los procesos de dispersión de semillas en la estructura de los bosques secos neotropicales. Uno de los principales hallazgos a partir de estos cuatro capítulos es que los patrones espaciales de las especies, así como las estrategias que utilizan para dispersarse y hacer frente a las condiciones adversas (es decir, lluvia o banco de semillas) llevan consigo un efecto del síndrome de dispersión, y que la intensidad ese efecto depende a la vez de las condiciones ambientales del lugar. ABSTRACT The importance of seed dispersal process in the estructuring and ecosystem dynamic is widely recongnized. However, for seasonally tropical dry forest studies related to this process are still scarce and scattered compared to tropical rain forests. The present research deals with the importance of seed dispersal syndromes as a driver in the community structure, focusing its attention to temporal and spatial patterns of seed dispersal in woody communities of seasonally dry forest at Southwestern Ecuador. This area is part of the Tumbesian region, one of the most important areas of endemism, but also one of the most threatened areas around the world. Climate is characterized by a dry season from May to November, and a rainy season from December to April. For the whole area an average temperature between 20 ° and 26 ° C, and an average annual rainfall between 300 and 700 mm are estimated. Fieldwork was carried out between February 2009 and September 2012. During a first step information about native woody species of dry forests of southwestern Ecuador was gathered, enabling to assign a dispersal syndrome to each species to determine the seed dispersal spectrum. In a second step, available information from 109 established plots along four municipalities in Loja province, which hold the highest and best conserved dry forest remanants, was analyzed to establish the relationship between dispersal syndromes and environmental conditions. The relationships between dispersal syndromes and species spatial patterns; and between dispersal syndromes and seed rain and seed bank patterns, were studied within a permanent plot of 9 ha, in the Arenillas Ecological Reserve. Within this plot one transect of approximately 3.4 km was set to collect monthly deer droppings, which were used to latter analyze the rol of this group as seed dispersers. The results showed that a large variety of plants in tropical dry forest of Southwestern Ecuador require animal assistance to dispers their seeds. However, an analysis of seed dispersal spectrum considering not only species richness, but also the relative abundance of species, allowed to determine that despite the high variety of zoochorous species, most individuals in the community corresponds to anemochoruos species. This shift may be due to the relative abundance of habitats that are suitable for species with different dispersal syndromes. Moreover, quantitative data analysis showed that environmental conditions affect the structure of seed dispersal spectrum in the studied community. The analysis of relative importance of dispersal syndrome, and the environmental heterogeneity on formation of adult trees spatial patterns, indicated that environmental heterogeneity exert an additional (or was the only) effect limiting the distribution of most species in this forest. The findings showed differences in spatial patterns related to dispersal syndrome, but also showed a large variation in spatial patterns even among species sharing the same dispersal syndrome. Simultaneous analysis of seed rain and seed bank patterns of a woody community, and their relationship with established vegetation, suggested that seed rain is temporally variable in species number and seeds abundance, and that variation is related to the dispersal syndrome. Dispersal syndrome also influenced on the formation of seed banks, being species with limited dispersal abilities (autochorous) the ones with highest species richness and seed abundance. Deer were found as a key element in the seed dispersal process. At least to eight woody species were dispersed legitimately by ingestion. Diaspores of most dispersed species had no obvious adaptations to seed dispersal, therefore, seed ingestion by deer represents a potential pathway for long-distance dispersal, and hence, improves the chances to colonizing new sites and to maintain gene flow. Overall, these results provide new evidence for understanding the importance of seed dispersal processes in the structure of Neotropical dry forests. One of the major findings from these four chapters is that spatial patterns of species, and the strategies used to disperse their seeds and to deal with the adverse conditions (i.e. seed rain or seed bank) are related with dispersal syndromes, and the intensity of that relation depends in turn, on environmental conditions.

Regenerative morphological traits in a woody species community in Tumbesian tropical dry forest.

The study of functional morphological traits enables us to know fundamental aspects of the dynamics of plant communities in local and global habitats. Regenerative morphological traits play an important role in defining plant history and ecological behavior. Seed and fruit characteristics determine to a large extent the patterns for dispersal, germination, establishment and seedling recruitment a given species exhibits on its natural habitat. Despite their prominent role, seed and fruit traits have been poorly studied at the community level of woody plant species in neo-tropical dry forests. In the present study we aimed at i) evaluate the functional role of morphological traits of seeds, fruits and embryo in woody plant species; ii) determine which are the morphological patterns present in seeds collected from the community of woody species that occur in neo-tropical dry forests; and iii) compare woody plant species seed mass values comparatively between neo-tropical dry and tropical forests. To do so, mature seeds were collected from 79 plant species that occur in the Tumbesian forest of Southwest Ecuador. The studied species included the 42 and 37 most representative tree and shrubbery species of the Tumbesian forest respectively. A total of 18 morphological traits (seven quantitative and 11 qualitative) were measured and evaluated in the seeds, fruits and embryos of the selected species, and we compared the seeds mass with other forest types. Our results showed a huge heterogeneity among traits values in the studied species. Seed mass, volume and number were the traits that vary the most at the community level, i.e. seed length ranged from 1.3 to 39 mm, and seed width from 0.6 to 25 mm. Only six embryo types were found among the 79 plant species. In 40 % of the cases, fully developed inverted embryos with large and thick cotyledons to store considerable amount of nutrients were recorded. We concluded that highly variable and functionally complementary morphological traits occur among the studied woody plants of the dry Tumbesian forest. The latter favors a plethora of behavioral mechanisms to coexist among woody species of the dry forest in response to the environmental stress that is typical of arid areas.

Seven Ulmus minor clones tolerant to Ophiostoma novo-ulmi registered as forest reproductive material in Spain

The Spanish elm programme began in 1986 in response to the devastating impact of Dutch elm disease on natural elm stands and urban trees. Its main objectives were to conserve remaining genetic resources and select and breed tolerant native elm genotypes. After 27 years of work conducting susceptibility trials on thousands of elm genotypes, the first seven tolerant Ulmus minor trees are now being registered by the Spanish Environmental Administration. This paper presents the results of the susceptibility tests on these clones and their distinctive genetic, morphological and phenological features. In all susceptibility trials the commercial ?Sapporo Autumn Gold? clone, which is highly tolerant to O. novo-ulmi, was used as a control. The registered clones were named ?Ademuz?, ?Dehesa de la Villa?, ?Majadahonda?, ?Toledo?, ?Dehesa de Amaniel?, ?Retiro? and ?Fuente Umbría?. The most tolerant clone was ?Dehesa de Amaniel?, as its wilting values were below 5% during the two consecutive inoculation trials performed in Madrid. ?Fuente Umbría?, tested over four consecutive years in Guadalajara and Palencia, was the Spanish clone with the most reliable tolerance level to O. novo-ulmi. The ?Ademuz? and ?Majadahonda? clones had the highest ornamental scores and are promising trees for use in urban environments and tree breeding for ornamental quality. These two genotypes showed a later bud burst phenology than the other U. minor clones, demonstrating suitability to areas with late frost events. The Spanish programme aims to substantially increase the range of tolerant native elms through new selections and crossings to gain a better understanding of the genetic basis of resistance.