Competition may explain the fine-scale spatial patterns and genetic structure of two co-occurring plant congeners

1. The spatial distribution of individual plants within a population and the population’s genetic structure are determined by several factors, like dispersal, reproduction mode or biotic interactions. The role of interspecific interactions in shaping the spatial genetic structure of plant populations remains largely unknown. 2. Species with a common evolutionary history are known to interact more closely with each other than unrelated species due to the greater number of traits they share. We hypothesize that plant interactions may shape the fine genetic structure of closely related congeners. 3. We used spatial statistics (georeferenced design) and molecular techniques (ISSR markers) to understand how two closely related congeners, Thymus vulgaris (widespread species) and T. loscosii (narrow endemic) interact at the local scale. Specific cover, number of individuals of both study species and several community attributes were measured in a 10 × 10 m plot. 4. Both species showed similar levels of genetic variation, but differed in their spatial genetic structure. Thymus vulgaris showed spatial aggregation but no spatial genetic structure, while T. loscosii showed spatial genetic structure (positive genetic autocorrelation) at short distances. The spatial pattern of T. vulgaris’ cover showed significant dissociation with that of T. loscosii. The same was true between the spatial patterns of the cover of T. vulgaris and the abundance of T. loscosii and between the abundance of each species. Most importantly, we found a correlation between the genetic structure of T. loscosii and the abundance of T. vulgaris: T. loscosii plants were genetically more similar when they were surrounded by a similar number of T. vulgaris plants. 5. Synthesis. Our results reveal spatially complex genetic structures of both congeners at small spatial scales. The negative association among the spatial patterns of the two species and the genetic structure found for T. loscosii in relation to the abundance of T. vulgaris indicate that competition between the two species may account for the presence of adapted ecotypes of T. loscosii to the abundance of a competing congeneric species. This suggests that the presence and abundance of close congeners can influence the genetic spatial structure of plant species at fine scales.

Spatial variability patterns of some Vertisol properties at a field scale using standardized data

Spatial variability of Vertisol properties is relevant for identifying those zones with physical degradation. In this sense, one has to face the problem of identifying the origin and distribution of spatial variability patterns. The objectives of the present work were (i) to quantify the spatial structure of different physical properties collected from a Vertisol, (ii) to search for potential correlations between different spatial patterns and (iii) to identify relevant components through multivariate spatial analysis. The study was conducted on a Vertisol (Typic Hapludert) dedicated to sugarcane (Saccharum officinarum L.) production during the last sixty years. We used six soil properties collected from a squared grid (225 points) (penetrometer resistance (PR), total porosity, fragmentation dimension (Df), vertical electrical conductivity (ECv), horizontal electrical conductivity (ECh) and soil water content (WC)). All the original data sets were z-transformed before geostatistical analysis. Three different types of semivariogram models were necessary for fitting individual experimental semivariograms. This suggests the different natures of spatial variability patterns. Soil water content rendered the largest nugget effect (C0 = 0.933) while soil total porosity showed the largest range of spatial correlation (A = 43.92 m). The bivariate geostatistical analysis also rendered significant cross-semivariance between different paired soil properties. However, four different semivariogram models were required in that case. This indicates an underlying co-regionalization between different soil properties, which is of interest for delineating management zones within sugarcane fields. Cross-semivariograms showed larger correlation ranges than individual, univariate, semivariograms (A ≥ 29 m). All the findings were supported by multivariate spatial analysis, which showed the influence of soil tillage operations, harvesting machinery and irrigation water distribution on the status of the investigated area.

Changes in spatial point patterns of pioneer woody plants across a large tropical landslide

We assessed whether the relative importance of positive and negative interactions in early successional communities varied across a large landslide on Casita Volcano (Nicaragua). We tested several hypotheses concerning the signatures of these processes in the spatial patterns of woody pioneer plants, as well as those of mortality and recruitment events, in several zones of the landslide differing in substrate stability and fertility, over a period of two years (2001 and 2002). We identified all woody individuals with a diameter >1 cm and mapped them in 28 plots measuring 10 × 10-m. On these maps, we performed a spatial point pattern analysis using univariate and bivariate pair-correlation functions; g (r) and g12 (r), and pairwise differences of univariate and bivariate functions. Spatial signatures of positive and negative interactions among woody plants were more prevalent in the most and least stressful zones of the landslide, respectively. Natural and human-induced disturbances such as the occurrence of fire, removal of newly colonizing plants through erosion and clearcutting of pioneer trees were also identified as potentially important pattern-creating processes. These results are in agreement with the stress-gradient hypothesis, which states that the relative importance of facilitation and competition varies inversely across gradients of abiotic stress. Our findings also indicate that the assembly of early successional plant communities in large heterogeneous landslides might be driven by a much larger array of processes than previously thought.

Lacunarity of the Spatial Distributions of Soil Types in Europe.

Lacunarity as a means of quantifying textural properties of spatial distributions suggests a classification into three main classes of the most abundant soils that cover 92% of Europe. Soils with a well-defined self-similar structure of the linear class are related to widespread spatial patterns that are nondominant but ubiquitous at continental scale. Fractal techniques have been increasingly and successfully applied to identify and describe spatial patterns in natural sciences. However, objects with the same fractal dimension can show very different optical properties because of their spatial arrangement. This work focuses primary attention on the geometrical structure of the geographical patterns of soils in Europe. We made use of the European Soil Database to estimate lacunarity indexes of the most abundant soils that cover 92% of the surface of Europe and investigated textural properties of their spatial distribution. We observed three main classes corresponding to three different patterns that displayed the graphs of lacunarity functions, that is, linear, convex, and mixed. They correspond respectively to homogeneous or self-similar, heterogeneous or clustered and those in which behavior can change at different ranges of scales. Finally, we discuss the pedological implications of that classification.

Mapping Genetic Diversity of Cherimoya (Annona cherimola Mill.): Application of Spatial Analysis for Conservation and Use of Plant Genetic Resources.

There is a growing call for inventories that evaluate geographic patterns in diversity of plant genetic resources maintained on farm and in species' natural populations in order to enhance their use and conservation. Such evaluations are relevant for useful tropical and subtropical tree species, as many of these species are still undomesticated, or in incipient stages of domestication and local populations can offer yet-unknown traits of high value to further domestication. For many outcrossing species, such as most trees, inbreeding depression can be an issue, and genetic diversity is important to sustain local production. Diversity is also crucial for species to adapt to environmental changes. This paper explores the possibilities of incorporating molecular marker data into Geographic Information Systems (GIS) to allow visualization and better understanding of spatial patterns of genetic diversity as a key input to optimize conservation and use of plant genetic resources, based on a case study of cherimoya (Annona cherimola Mill.), a Neotropical fruit tree species. We present spatial analyses to (1) improve the understanding of spatial distribution of genetic diversity of cherimoya natural stands and cultivated trees in Ecuador, Bolivia and Peru based on microsatellite molecular markers (SSRs); and (2) formulate optimal conservation strategies by revealing priority areas for in situ conservation, and identifying existing diversity gaps in ex situ collections. We found high levels of allelic richness, locally common alleles and expected heterozygosity in cherimoya's putative centre of origin, southern Ecuador and northern Peru, whereas levels of diversity in southern Peru and especially in Bolivia were significantly lower. The application of GIS on a large microsatellite dataset allows a more detailed prioritization of areas for in situ conservation and targeted collection across the Andean distribution range of cherimoya than previous studies could do, i.e. at province and department level in Ecuador and Peru, respectively.

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

Juvenile–adult tree associations in a continental Mediterranean ecosystem: no evidence for sustained and general facilitation at increased aridity

Question: How do tree species identity, microhabitat and water availability affect inter- and intra-specific interactions between juvenile and adult woody plants? Location: Continental Mediterranean forests in Alto Tajo Natural Park, Guadalajara, Spain. Methods: A total of 2066 juveniles and adults of four co-occurring tree species were mapped in 17 plots. The frequency of juveniles at different microhabitats and water availability levels was analysed using log-linear models. We used nearest-neighbour contingency table analysis of spatial segregation and J-functions to describe the spatial patterns. Results: We found a complex spatial pattern that varied according to species identity and microhabitat. Recruitment was more frequent in gaps for Quercus ilex, while the other three species recruited preferentially under shrubs or trees depending on the water availability level. Juveniles were not spatially associated to conspecific adults, experiencing segregation from them inmany cases. Spatial associations, both positive and negative, were more common at higher water availability levels. Conclusions: Our results do not agree with expectations from the stressgradient hypothesis, suggesting that positive interactions do not increase in importance with increasing aridity in the study ecosystem. Regeneration patterns are species-specific and depend on microhabitat characteristics and dispersal strategies. In general, juveniles do not look for conspecific adult protection. This work contributes to the understanding of species co-existence, proving the importance of considering a multispecies approach at several plots to overcome limitations of simple pair-wise comparisons in a limited number of sites.

Flying vs. climbing: Factors controlling arboreal seed removal in oak-beech forests

Nuts are heavy and nutritious seeds that need animals to be successfully dispersed. Most studies address nut removal by a single animal species once seeds fall onto the ground. However, nuts are also accessible before the seed drop and usually to a wide guild of seed foragers. This study examines the factorscontrollingarborealseedremoval in oak–beechforests within the whole guild of nut foragers. We found that seed-dispersing rodents (Apodemus sylvaticus) were the main acorn removers in the oaks (up to 3.75 m height), with a rapid seed encounter and a high removal rate. However, rodents did not climb the beech trees, probably due to their smoother bark in comparison to oak bark and/or the lower nutritional value of beechnuts with regard to acorns. Jays (Garrulus glandarius) were more abundant in oak stands (both dense and scattered) and clearly preferred acorns to beechnuts whereas nuthatches (Sitta europaea) were more abundant in beech stands and preferred beechnuts to acorns. Non-storing birds such as great tits (Parus major) also removed acorns and beechnuts, especially in the stands where oaks are dominant. Jays and rodents preferred sound seeds over insect-infested seeds but such a preference was not found for nuthatches. This study highlights that pure beech stands showed a reduced guild of arboreal nut foragers in comparison to oak stands. This different guild could probably affect the spatial patterns of seed dispersal, with a proportionally higher number of long dispersal events for acorns (mostly jay-dispersed) than for beechnuts (mostly nuthatch-dispersed). Long-distance dispersal of beechnuts (by jays) is determined by the presence of other preferred species (oaks) and their frequency of non-mast years. Seed location in different habitats strongly determines the contribution of different arboreal removers (including climbing rodents) and their removal speed, leading to a differential seed fate that will eventually affect tree regeneration. As nuthatches are sedentary birds, it is important to maintain old and dead trees where they can breed (crevices), forage (arthropods) and store seeds in order to favor beechnut dispersal and gene flow. By maintaining or favoring oak trees within beech stands we will ensure a wider guild of arboreal nut dispersers.

Fractal scaling of apparent soil moisture estimated from vertical planes of Vertisol pit images

Image analysis could be a useful tool for investigating the spatial patterns of apparent soil moisture at multiple resolutions. The objectives of the present work were (i) to define apparent soil moisture patterns from vertical planes of Vertisol pit images and (ii) to describe the scaling of apparent soil moisture distribution using fractal parameters.

Fractal scaling of apparent soil moisture estimated from vertical planes of Vertisol pit images

Image analysis could be a useful tool for investigating the spatial patterns of apparent soil moisture at multiple resolutions. The objectives of the present work were (i) to define apparent soil moisture patterns from vertical planes of Vertisol pit images and (ii) to describe the scaling of apparent soil moisture distribution using fractal parameters. Twelve soil pits (0.70 m long × 0.60 m width × 0.30 m depth) were excavated on a bare Mazic Pellic Vertisol. Six of them were excavated in April/2011 and six pits were established in May/2011 after 3 days of a moderate rainfall event. Digital photographs were taken from each Vertisol pit using a Kodak™ digital camera. The mean image size was 1600 × 945 pixels with one physical pixel ≈373 μm of the photographed soil pit. Each soil image was analyzed using two fractal scaling exponents, box counting (capacity) dimension (DBC) and interface fractal dimension (Di), and three prefractal scaling coefficients, the total number of boxes intercepting the foreground pattern at a unit scale (A), fractal lacunarity at the unit scale (Λ1) and Shannon entropy at the unit scale (S1). All the scaling parameters identified significant differences between both sets of spatial patterns. Fractal lacunarity was the best discriminator between apparent soil moisture patterns. Soil image interpretation with fractal exponents and prefractal coefficients can be incorporated within a site-specific agriculture toolbox. While fractal exponents convey information on space filling characteristics of the pattern, prefractal coefficients represent the investigated soil property as seen through a higher resolution microscope. In spite of some computational and practical limitations, image analysis of apparent soil moisture patterns could be used in connection with traditional soil moisture sampling, which always renders punctual estimates

Estrategias innovadoras para la gestión del riesgo de sequía en sistemas ganaderos extensivos

La vulnerabilidad de los sistemas ganaderos de pastoreo pone en evidencia la necesidad de herramientas para evaluar y mitigar los efectos de la sequía. El avance en la teledetección ha despertado el interés por explotar potenciales aplicaciones, y está dando lugar a un intenso desarrollo de innovaciones en distintos campos. Una de estas áreas es la gestión del riesgo climático, en donde la utilización de índices de vegetación permite la evaluación de la sequía. En esta investigación, se analiza el impacto de la sequía y se evalúa el potencial de nuevas tecnologías como la teledetección para la gestión del riesgo de sequía en sistemas de ganadería extensiva. Para ello, se desarrollan tres aplicaciones: (i) evaluar el impacto económico de la sequía en una explotación ganadera extensiva de la dehesa de Andalucía, (ii) elaborar mapas de vulnerabilidad a la sequía en pastos de Chile y (iii) diseñar y evaluar el potencial de un seguro indexado para sequía en pastos en la región de Coquimbo en Chile. En la primera aplicación, se diseña un modelo dinámico y estocástico que integra aspectos climáticos, ecológicos, agronómicos y socioeconómicos para evaluar el riesgo de sequía. El modelo simula una explotación ganadera tipo de la dehesa de Andalucía para el período 1999-2010. El método de Análisis Histórico y la simulación de MonteCarlo se utilizan para identificar los principales factores de riesgo de la explotación, entre los que destacan, los periodos de inicios del verano e inicios de invierno. Los resultados muestran la existencia de un desfase temporal entre el riesgo climático y riesgo económico, teniendo este último un periodo de duración más extenso en el tiempo. También, revelan que la intensidad, frecuencia y duración son tres atributos cruciales que determinan el impacto económico de la sequía. La estrategia de reducción de la carga ganadera permite aminorar el riesgo, pero conlleva una disminución en el margen bruto de la explotación. La segunda aplicación está dedicada a la elaboración de mapas de vulnerabilidad a la sequia en pastos de Chile. Para ello, se propone y desarrolla un índice de riesgo económico (IRESP) sencillo de interpretar y replicable, que integra factores de riesgo y estrategias de adaptación para obtener una medida del Valor en Riesgo, es decir, la máxima pérdida esperada en un año con un nivel de significación del 5%.La representación espacial del IRESP pone en evidencia patrones espaciales y diferencias significativas en la vulnerabilidad a la sequía a lo largo de Chile. Además, refleja que la vulnerabilidad no siempre esta correlacionada con el riesgo climático y demuestra la importancia de considerar las estrategias de adaptación. Las medidas de autocorrelación espacial revelan que el riesgo sistémico es considerablemente mayor en el sur que en el resto de zonas. Los resultados demuestran que el IRESP transmite información pertinente y, que los mapas de vulnerabilidad pueden ser una herramienta útil en el diseño de políticas y toma de decisiones para la gestión del riesgo de sequía. La tercera aplicación evalúa el potencial de un seguro indexado para sequía en pastos en la región de Coquimbo en Chile. Para lo cual, se desarrolla un modelo estocástico para estimar la prima actuarialmente justa del seguro y se proponen y evalúan pautas alternativas para mejorar el diseño del contrato. Se aborda el riesgo base, el principal problema de los seguros indexados identificado en la literatura y, que está referido a la correlación imperfecta del índice con las pérdidas de la explotación. Para ello, se sigue un enfoque bayesiano que permite evaluar el impacto en el riesgo base de las pautas de diseño propuestas: i) una zonificación por clúster que considera aspectos espacio-temporales, ii) un período de garantía acotado a los ciclos fenológicos del pasto y iii) umbral de garantía. Los resultados muestran que tanto la zonificación como el periodo de garantía reducen el riesgo base considerablemente. Sin embargo, el umbral de garantía tiene un efecto ambiguo sobre el riesgo base. Por otra parte, la zonificación por clúster contribuye a aminorar el riesgo sistémico que enfrentan las aseguradoras. Estos resultados han puesto de manifiesto que un buen diseño de contrato puede tener un doble dividendo, por un lado aumentar su utilidad y, por otro, reducir el coste del seguro. Un diseño de contrato eficiente junto con los avances en la teledetección y un adecuado marco institucional son los pilares básicos para el buen funcionamiento de un programa de seguro. Las nuevas tecnologías ofrecen un importante potencial para la innovación en la gestión del riesgo climático. Los avances en este campo pueden proporcionar importantes beneficios sociales en los países en desarrollo y regiones vulnerables, donde las herramientas para gestionar eficazmente los riesgos sistémicos como la sequía pueden ser de gran ayuda para el desarrollo. The vulnerability of grazing livestock systems highlights the need for tools to assess and mitigate the adverse impact of drought. The recent and rapid progress in remote sensing has awakened an interest for tapping into potential applications, triggering intensive efforts to develop innovations in a number of spheres. One of these areas is climate risk management, where the use of vegetation indices facilitates assessment of drought. This research analyzes drought impacts and evaluates the potential of new technologies such as remote sensing to manage drought risk in extensive livestock systems. Three essays in drought risk management are developed to: (i) assess the economic impact of drought on a livestock farm in the Andalusian Dehesa, (ii) build drought vulnerability maps in Chilean grazing lands, and (iii) design and evaluate the potential of an index insurance policy to address the risk of drought in grazing lands in Coquimbo, Chile. In the first essay, a dynamic and stochastic farm model is designed combining climate, agronomic, socio-economic and ecological aspects to assess drought risk. The model is developed to simulate a representative livestock farm in the Dehesa of Andalusia for the time period 1999-2010. Burn analysis and MonteCarlo simulation methods are used to identify the significance of various risk sources at the farm. Most notably, early summer and early winter are identified as periods of peak risk. Moreover, there is a significant time lag between climate and economic risk and this later last longer than the former. It is shown that intensity, frequency and duration of the drought are three crucial attributes that shape the economic impact of drought. Sensitivity analysis is conducted to assess the sustainability of farm management strategies and demonstrates that lowering the stocking rate reduces farmer exposure to drought risk but entails a reduction in the expected gross margin. The second essay, mapping drought vulnerability in Chilean grazing lands, proposes and builds an index of economic risk (IRESP) that is replicable and simple to interpret. This methodology integrates risk factors and adaptation strategies to deliver information on Value at Risk, maximum expected losses at 5% significance level. Mapping IRESP provides evidence about spatial patterns and significant differences in drought vulnerability across Chilean grazing lands. Spatial autocorrelation measures reveal that systemic risk is considerably larger in the South as compared to Northern or Central Regions. Furthermore, it is shown that vulnerability is not necessarily correlated with climate risk and that adaptation strategies do matter. These results show that IRESP conveys relevant information and that vulnerability maps may be useful tools to assess policy design and decision-making in drought risk management. The third essay develops a stochastic model to estimate the actuarially fair premium and evaluates the potential of an indexed insurance policy to manage drought risk in Coquimbo, a relevant livestock farming region of Chile. Basis risk refers to the imperfect correlation of the index and farmer loses and is identified in the literature as a main limitation of index insurance. A Bayesian approach is proposed to assess the impact on basis risk of alternative guidelines in contract design: i) A cluster zoning that considers space-time aspects, ii) A guarantee period bounded to fit phenological cycles, and iii) the triggering index threshold. Results show that both the proposed zoning and guarantee period considerably reduces basis risk. However, the triggering index threshold has an ambiguous effect on basis risk. On the other hand, cluster zoning contributes to ameliorate systemic risk faced by the insurer. These results highlighted that adequate contract design is important and may result in double dividend. On the one hand, increasing farmers’ utility and, secondly, reducing the cost of insurance. An efficient contract design coupled with advances in remote sensing and an appropriate institutional framework are the basis for an efficient operation of an insurance program. The new technologies offer significant potential for innovation in climate risk managements. Progress in this field is capturing increasing attention and may provide important social gains in developing countries and vulnerable regions where the tools to efficiently manage systemic risks, such as drought, may be a means to foster development.

Fractal scaling of apparent soil moisture estimated from vertical planesof Vertisol pit images

Image analysis could be a useful tool for investigating the spatial patterns of apparent soil moisture at multiple resolutions. The objectives of the present work were (i) to define apparent soil moisture patterns from vertical planes of Vertisol pit images and (ii) to describe the scaling of apparent soil moisture distribution using fractal parameters. Twelve soil pits (0.70 m long × 0.60 m width × 0.30 m depth) were excavated on a bare Mazic Pellic Vertisol. Six of them were excavated in April/2011 and six pits were established in May/2011 after 3 days of a moderate rainfall event. Digital photographs were taken from each Vertisol pit using a Kodak? digital camera. The mean image size was 1600 × 945 pixels with one physical pixel ?373 ?m of the photographed soil pit. Each soil image was analyzed using two fractal scaling exponents, box counting (capacity) dimension (DBC) and interface fractal dimension (Di), and three prefractal scaling coefficients, the total number of boxes intercepting the foreground pattern at a unit scale (A), fractal lacunarity at the unit scale (?1) and Shannon entropy at the unit scale (S1). All the scaling parameters identified significant differences between both sets of spatial patterns. Fractal lacunarity was the best discriminator between apparent soil moisture patterns. Soil image interpretation with fractal exponents and prefractal coefficients can be incorporated within a site-specific agriculture toolbox. While fractal exponents convey information on space filling characteristics of the pattern, prefractal coefficients represent the investigated soil property as seen through a higher resolution microscope. In spite of some computational and practical limitations, image analysis of apparent soil moisture patterns could be used in connection with traditional soil moisture sampling, which always renders punctual estimates.

Implementation of a module for risk of ozone impacts assessment to vegetation in the integrated assessment modelling system for the Iberian peninsula. Evaluation for wheat and holm oak

A module to estimate risks of ozone damage to vegetation has been implemented in the Integrated Assessment Modelling system for the Iberian Peninsula. It was applied to compute three different indexes for wheat and Holm oak; daylight AOT40 (cumulative ozone concentration over 40 ppb), cumulative ozone exposure index according to the Directive 2008/50/EC (AOT40-D) and PODY (Phytotoxic Ozone Dose over a given threshold of Y nmol m−2 s−1). The use of these indexes led to remarkable differences in spatial patterns of relative ozone risks on vegetation. Ozone critical levels were exceeded in most of the modelling domain and soil moisture content was found to have a significant impact on the results. According to the outputs of the model, daylight AOT40 constitutes a more conservative index than the AOT40-D. Additionally, flux-based estimations indicate high risk areas in Portugal for both wheat and Holm oak that are not identified by AOT-based methods.

Toward a climate-dependent paradigm of ammonia emission & deposition.

Existing descriptions of bi-directional ammonia (NH3) land–atmosphere exchange incorporate temperature and moisture controls, and are beginning to be used in regional chemical transport models. However, such models have typically applied simpler emission factors to upscale the main NH3 emission terms. While this approach has successfully simulated the main spatial patterns on local to global scales, it fails to address the environment- and climate-dependence of emissions. To handle these issues, we outline the basis for a new modelling paradigm where both NH3 emissions and deposition are calculated online according to diurnal, seasonal and spatial differences in meteorology. We show how measurements reveal a strong, but complex pattern of climatic dependence, which is increasingly being characterized using ground-based NH3 monitoring and satellite observations, while advances in process-based modelling are illustrated for agricultural and natural sources, including a global application for seabird colonies. A future architecture for NH3 emission–deposition modelling is proposed that integrates the spatio-temporal interactions, and provides the necessary foundation to assess the consequences of climate change. Based on available measurements, a first empirical estimate suggests that 5°C warming would increase emissions by 42 per cent (28–67%). Together with increased anthropogenic activity, global NH3 emissions may increase from 65 (45–85) Tg N in 2008 to reach 132 (89–179) Tg by 2100.

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.