Seed Dispersal Spectrum of Woody Species in South Ecuadorian Dry Forests: Environmental Correlates and the Effect of Considering Species Abundance

This study examines the seed dispersal spectrum of the tropical dry forests of Southern Ecuador, in an effort to contribute to the knowledge of the complex dynamics of tropical dry forests. Seed dispersal spectrum was described for a total number of 160 species. Relationships of dispersal syndromes with plant growth form and climatic seasonality were explored. For a subset of 97 species, we determined whether dispersal spectrum changes when species abundance, in addition to species number, is taken into account. The same subset was used to relate dispersal syndromes with the environmental conditions. Zoochorous species dominated in the studied community. When considering the individual abundance of each species, however, anemochory was the prevalent dispersal syndrome. We found a significant difference in the frequency of dispersal syndromes among plant growth forms, with epizoochory only occurring in shrub species. The dispersal spectrum was dependent on climatic seasonality. The largest proportion of anemochorous species fructified during the dry season, while zoochorous diaspores dominated during the rainy season. A fourth-corner analysis indicated that the seed dispersal spectrum of Southern Ecuador dry forests is controlled by environmental conditions such as annual precipitation, annual temperature range or topography. Our results suggest that spatio-temporal changes in the environmental conditions may affect important ecological processes for dispersal. Thus, the predominance of one syndrome or another may depend on the spatial variation of environmental conditions.

Climatic factors control rodent seed predation in Pinus pinea L. stands in Central Spain

- Context: Pinus pinea L. presents serious problems of natural regeneration in managed forest of Central Spain. The species exhibits specific traits linked to frugivore activity. Therefore, information on plant–animal interactions may be crucial to understand regeneration failure. - Aims: Determining the spatio-temporal pattern of P. pinea seed predation by Apodemus sylvaticus L. and the factors involved. Exploring the importance of A. sylvaticus L. as a disperser of P. pinea. Identifying other frugivores and their seasonal patterns. - Methods: An intensive 24-month seed predation trial was carried out. The probability of seeds escaping predation was modelled through a zero-inflated binomial mixed model. Experiments on seed dispersal by A. sylvaticus were conducted. Cameras were set up to identify other potential frugivores. - Results: Decreasing rodent population in summer and masting enhances seed survival. Seeds were exploited more rapidly nearby parent trees and shelters. A. sylvaticus dispersal activity was found to be scarce. Corvids marginally preyed upon P. pinea seeds. - Conclusions: Survival of P. pinea seeds is climate-controlled through the timing of the dry period together with masting occurrence. Should germination not take place during the survival period, establishment may be limited. A. sylvaticus mediated dispersal does not modify the seed shadow. Seasonality of corvid activity points to a role of corvids in dispersal.

Trends in climatic variables and future reference evapotranspiration in Duero Valley (Spain)

The impact of climate change and its relation with evapotranspiration was evaluated in the Duero River Basin (Spain). The study shows possible future situations 50 yr from now from the reference evapotranspiration (ETo). The maximum temperature (Tmax), minimum temperature (Tmin), dew point (Td), wind speed (U) and net radiation (Rn) trends during the 1980–2009 period were obtained and extrapolated with the FAO-56 Penman-Montheith equation to estimate ETo. Changes in stomatal resistance in response to increases in CO2 were also considered. Four scenarios were done, taking the concentration of CO2 and the period analyzed (annual or monthly) into consideration. The scenarios studied showed the changes in ETo as a consequence of the annual and monthly trends in the variables Tmax, Tmin, Td, U and Rn with current and future CO2 concentrations (372 ppm and 550 ppm). The future ETo showed increases between 118 mm (11 %) and 55 mm (5 %) with respect to the current situation of the river basin at 1042 mm. The months most affected by climate change are May, June, July, August and September, which also coincide with the maximum water needs of the basin’s crops

Levels, sources and seasonality of coarse particles (PM10-PM2.5) in three European capitals e implications for particulate pollution control

Coarse particles of aerodynamic diameter between 2.5 and 10 mm (PMc) are produced by a range of natural (windblown dust and sea sprays) and anthropogenic processes (non-exhaust vehicle emissions, industrial, agriculture, construction and quarrying activities). Although current ambient air quality regulations focus on PM2.5 and PM10, coarse particles are of interest from a public health point of view as they have been associated with certain mortality and morbidity outcomes. In this paper, an analysis of coarse particle levels in three European capitals (London, Madrid and Athens) is presented and discussed. For all three cities we analysed data from both traffic and urban background monitoring sites. The results showed that the levels of coarse particles present significant seasonal, weekly and daily variability. Their wind driven and non-wind driven resuspension as well as their roadside increment due to traffic were estimated. Both the local meteorological conditions and the air mass history indicating long-range atmospheric transport of particles of natural origin are significant parameters that influence the levels of coarse particles in the three cities especially during episodic events.

Extreme pointer years in tree-ring records of central Spain as evidence of climatic events and the eruption of the Huaynaputina Volcano (Peru, 1600 AD)

The study of pointer years of numerous tree-ring chronologies of the central Iberian Peninsula (Sierra de Guadarrama) could provide complementary information about climate variability over the last 405 yr. In total, 64 pointer years have been identified: 30 negative (representing minimum growths) and 34 positive (representing maximum growths), the most significant of these being 1601, 1963 and 1996 for the negative ones, and 1734 and 1737 for the positive ones. Given that summer precipitation was found to be the most limiting factor for the growth of Pinus in the Sierra de Guadarrama in the second half of the 20th century, it is also an explanatory factor in almost 50% of the extreme growths. Furthermore, these pointer years and intervals are not evenly distributed throughout time. Both in the first half of the 17th and in the second half of 20th, they were more frequent and more extreme and these periods are the most notable for the frequency of negative pointer years in Central Spain. The interval 1600–1602 is of special significance, being one of the most unfavourable for tree growth in the centre of Spain, with 1601 representing the minimum index in the regional chronology. We infer that this special minimum annual increase was the effect of the eruption of Huaynaputina, which occurred in Peru at the beginning of 1600 AD. This is the first time that the effects of this eruption in the tree-ring records of Southern Europe have been demonstrated.

Modelling Pinus pinea forest management to attain natural regeneration under present and future climatic scenarios

Natural regeneration-based silviculture has been increasingly regarded as a reliable option in sustainable forest management. However, successful natural regeneration is not always easy to achieve. Recently, new concerns have arisen because of changing future climate. To date, regeneration models have proved helpful in decision-making concerning natural regeneration. The implementation of such models into optimization routines is a promising approach in providing forest managers with accurate tools for forest planning. In the present study, we present a stochastic multistage regeneration model for Pinus pinea L. managed woodlands in Central Spain, where regeneration has been historically unsuccessful. The model is able to quantify recruitment under different silviculture alternatives and varying climatic scenarios, with further application to optimize management scheduling. The regeneration process in the species showed high between-year variation, with all subprocesses (seed production, dispersal, germination, predation, and seedling survival) having the potential to become bottlenecks. However, model simulations demonstrate that current intensive management is responsible for regeneration failure in the long term. Specifically, stand densities at rotation age are too low to guarantee adequate dispersal, the optimal density of seed-producing trees being around 150 stems·ha−1. In addition, rotation length needs to be extended up to 120 years to benefit from the higher seed production of older trees. Stochastic optimization confirms these results. Regeneration does not appear to worsen under climate change conditions; the species exhibiting resilience worthy of broader consideration in Mediterranean silviculture.

Carbon allocation in north-western Amazon forests (Colombia)

Los estudios sobre la asignación del carbono en los ecosistemas forestales proporcionan información esencial para la comprensión de las diferencias espaciales y temporales en el ciclo del carbono de tal forma que pueden aportar información a los modelos y, así predecir las posibles respuestas de los bosques a los cambios en el clima. Dentro de este contexto, los bosques Amazónicos desempeñan un papel particularmente importante en el balance global del carbono; no obstante, existen grandes incertidumbres en cuanto a los controles abióticos en las tasas de la producción primaria neta (PPN), la asignación de los productos de la fotosíntesis a los diferentes componentes o compartimentos del ecosistema (aéreo y subterráneo) y, cómo estos componentes de la asignación del carbono responden a eventos climáticos extremos. El objetivo general de esta tesis es analizar los componentes de la asignación del carbono en bosques tropicales maduros sobre suelos contrastantes, que crecen bajo condiciones climáticas similares en dos sitios ubicados en la Amazonia noroccidental (Colombia): el Parque Natural Nacional Amacayacu y la Estación Biológica Zafire. Con este objetivo, realicé mediciones de los componentes de la asignación del carbono (biomasa, productividad primaria neta, y su fraccionamiento) a nivel ecosistémico y de la dinámica forestal (tasas anuales de mortalidad y reclutamiento), a lo largo de ocho años (20042012) en seis parcelas permanentes de 1 hectárea establecidas en cinco tipos de bosques sobre suelos diferentes (arcilloso, franco-arcilloso, franco-arcilloso-arenoso, franco-arenoso y arena-francosa). Toda esta información me permitió abordar preguntas específicas que detallo a continuación. En el Capítulo 2 evalúe la hipótesis de que a medida que aumenta la fertilidad del suelo disminuye la cantidad del carbono asignado a la producción subterránea (raíces finas con diámetro <2 mm). Y para esto, realicé mediciones de la masa y la producción de raíces finas usando dos métodos: (1) el de los cilindros de crecimiento y, (2) el de los cilindros de extracción secuencial. El monitoreo se realizó durante 2.2 años en los bosques con suelos más contrastantes: arcilla y arena-francosa. Encontré diferencias significativas en la masa de raíces finas y su producción entre los bosques y, también con respecto a la profundidad del suelo (010 y 1020 cm). El bosque sobre arena-francosa asignó más carbono a las raíces finas que el bosque sobre arcillas. La producción de raíces finas en el bosque sobre arena-francosa fue dos veces más alta (media ± error estándar = 2.98 ± 0.36 y 3.33 ± 0.69 Mg C ha1 año1, con el método 1 y 2, respectivamente), que para el bosque sobre arcillas, el suelo más fértil (1.51 ± 0.14, método 1, y desde 1.03 ± 0.31 a 1.36 ± 0.23 Mg C ha1 año1, método 2). Del mismo modo, el promedio de la masa de raíces finas fue tres veces mayor en el bosque sobre arena-francosa (5.47 ± 0.17 Mg C ha1) que en el suelo más fértil (de 1.52 ± 0.08 a 1.82 ± 0.09 Mg C ha1). La masa de las raíces finas también mostró un patrón temporal relacionado con la lluvia, mostrando que la producción de raíces finas disminuyó sustancialmente en el período seco del año 2005. Estos resultados sugieren que los recursos del suelo pueden desempeñar un papel importante en los patrones de la asignación del carbono entre los componentes aéreo y subterráneo de los bosques tropicales; y que el suelo no sólo influye en las diferencias en la masa de raíces finas y su producción, sino que también, en conjunto con la lluvia, sobre la estacionalidad de la producción. En el Capítulo 3 estimé y analicé los tres componentes de la asignación del carbono a nivel del ecosistema: la biomasa, la productividad primaria neta PPN, y su fraccionamiento, en los mismos bosques del Capítulo 2 (el bosque sobre arcillas y el bosque sobre arena-francosa). Encontré diferencias significativas en los patrones de la asignación del carbono entre los bosques; el bosque sobre arcillas presentó una mayor biomasa total y aérea, así como una PPN, que el bosque sobre arena-francosa. Sin embargo, la diferencia entre los dos bosques en términos de la productividad primaria neta total fue menor en comparación con las diferencias entre la biomasa total de los bosques, como consecuencia de las diferentes estrategias en la asignación del carbono a los componentes aéreo y subterráneo del bosque. La proporción o fracción de la PPN asignada a la nueva producción de follaje fue relativamente similar entre los dos bosques. Nuestros resultados de los incrementos de la biomasa aérea sugieren una posible compensación entre la asignación del carbono al crecimiento de las raíces finas versus el de la madera, a diferencia de la compensación comúnmente asumida entre la parte aérea y la subterránea en general. A pesar de estas diferencias entre los bosques en términos de los componentes de la asignación del carbono, el índice de área foliar fue relativamente similar entre ellos, lo que sugiere que el índice de área foliar es más un indicador de la PPN total que de la asignación de carbono entre componentes. En el Capítulo 4 evalué la variación espacial y temporal de los componentes de la asignación del carbono y la dinámica forestal de cinco tipos e bosques amazónicos y sus respuestas a fluctuaciones en la precipitación, lo cual es completamente relevante en el ciclo global del carbono y los procesos biogeoquímicos en general. Estas variaciones son así mismo importantes para evaluar los efectos de la sequía o eventos extremos sobre la dinámica natural de los bosques amazónicos. Evalué la variación interanual y la estacionalidad de los componentes de la asignación del carbono y la dinámica forestal durante el periodo 2004−2012, en cinco bosques maduros sobre diferentes suelos (arcilloso, franco-arcilloso, franco-arcilloso-arenoso, franco-arenoso y arena-francosa), todos bajo el mismo régimen local de precipitación en la Amazonia noroccidental (Colombia). Quería examinar sí estos bosques responden de forma similar a las fluctuaciones en la precipitación, tal y como pronostican muchos modelos. Consideré las siguientes preguntas: (i) ¿Existe una correlación entre los componentes de la asignación del carbono y la dinámica forestal con la precipitación? (ii) ¿Existe correlación entre los bosques? (iii) ¿Es el índice de área foliar (LAI) un indicador de las variaciones en la producción aérea o es un reflejo de los cambios en los patrones de la asignación del carbono entre bosques?. En general, la correlación entre los componentes aéreo y subterráneo de la asignación del carbono con la precipitación sugiere que los suelos juegan un papel importante en las diferencias espaciales y temporales de las respuestas de estos bosques a las variaciones en la precipitación. Por un lado, la mayoría de los bosques mostraron que los componentes aéreos de la asignación del carbono son susceptibles a las fluctuaciones en la precipitación; sin embargo, el bosque sobre arena-francosa solamente presentó correlación con la lluvia con el componente subterráneo (raíces finas). Por otra parte, a pesar de que el noroeste Amazónico es considerado sin una estación seca propiamente (definida como <100 mm meses −1), la hojarasca y la masa de raíces finas mostraron una alta variabilidad y estacionalidad, especialmente marcada durante la sequía del 2005. Además, los bosques del grupo de suelos francos mostraron que la hojarasca responde a retrasos en la precipitación, al igual que la masa de raíces finas del bosque sobre arena-francosa. En cuanto a la dinámica forestal, sólo la tasa de mortalidad del bosque sobre arena-francosa estuvo correlacionada con la precipitación (ρ = 0.77, P <0.1). La variabilidad interanual en los incrementos en el tallo y la biomasa de los individuos resalta la importancia de la mortalidad en la variación de los incrementos en la biomasa aérea. Sin embargo, las tasas de mortalidad y las proporciones de individuos muertos por categoría de muerte (en pie, caído de raíz, partido y desaparecido), no mostraron tendencias claras relacionadas con la sequía. Curiosamente, la hojarasca, el incremento en la biomasa aérea y las tasas de reclutamiento mostraron una alta correlación entre los bosques, en particular dentro del grupo de los bosques con suelos francos. Sin embargo, el índice de área foliar estimado para los bosques con suelos más contrastantes (arcilla y arena-francosa), no presentó correlación significativa con la lluvia; no obstante, estuvo muy correlacionado entre bosques; índice de área foliar no reflejó las diferencias en la asignación de los componentes del carbono, y su respuesta a la precipitación en estos bosques. Por último, los bosques estudiados muestran que el noroeste amazónico es susceptible a fenómenos climáticos, contrario a lo propuesto anteriormente debido a la ausencia de una estación seca propiamente dicha. ABSTRACT Studies of carbon allocation in forests provide essential information for understanding spatial and temporal differences in carbon cycling that can inform models and predict possible responses to changes in climate. Amazon forests play a particularly significant role in the global carbon balance, but there are still large uncertainties regarding abiotic controls on the rates of net primary production (NPP) and the allocation of photosynthetic products to different ecosystem components; and how the carbon allocation components of Amazon forests respond to extreme climate events. The overall objective of this thesis is to examine the carbon allocation components in old-growth tropical forests on contrasting soils, and under similar climatic conditions in two sites at the Amacayacu National Natural Park and the Zafire Biological Station, located in the north-western Amazon (Colombia). Measurements of above- and below-ground carbon allocation components (biomass, net primary production, and its partitioning) at the ecosystem level, and dynamics of tree mortality and recruitment were done along eight years (20042012) in six 1-ha plots established in five Amazon forest types on different soils (clay, clay-loam, sandy-clay-loam, sandy-loam and loamy-sand) to address specific questions detailed in the next paragraphs. In Chapter 2, I evaluated the hypothesis that as soil fertility increases the amount of carbon allocated to below-ground production (fine-roots) should decrease. To address this hypothesis the standing crop mass and production of fine-roots (<2 mm) were estimated by two methods: (1) ingrowth cores and, (2) sequential soil coring, during 2.2 years in the most contrasting forests: the clay-soil forest and the loamy-sand forest. We found that the standing crop fine-root mass and its production were significantly different between forests and also between soil depths (0–10 and 10–20 cm). The loamysand forest allocated more carbon to fine-roots than the clay-soil forest, with fine-root production in the loamy-sand forest twice (mean ± standard error = 2.98 ± 0.36 and 3.33 ± 0.69 Mg C ha −1 yr −1, method 1 and 2, respectively) as much as for the more fertile claysoil forest (1.51 ± 0.14, method 1, and from 1.03 ± 0.31 to 1.36 ± 0.23 Mg C ha −1 yr −1, method 2). Similarly, the average of standing crop fine-root mass was three times higher in the loamy-sand forest (5.47 ± 0.17 Mg C ha1) than in the more fertile soil (from 1.52 ± 0.08 a 1.82 ± 0.09 Mg C ha1). The standing crop fine-root mass also showed a temporal pattern related to rainfall, with the production of fine-roots decreasing substantially in the dry period of the year 2005. These results suggest that soil resources may play an important role in patterns of carbon allocation of below-ground components, not only driven the differences in the biomass and its production, but also in the time when it is produced. In Chapter 3, I assessed the three components of stand-level carbon allocation (biomass, NPP, and its partitioning) for the same forests evaluated in Chapter 2 (clay-soil forest and loamy-sand forest). We found differences in carbon allocation patterns between these two forests, showing that the forest on clay-soil had a higher aboveground and total biomass as well as a higher above-ground NPP than the loamy-sand forest. However, differences between the two types of forests in terms of stand-level NPP were smaller, as a consequence of different strategies in the carbon allocation of above- and below-ground components. The proportional allocation of NPP to new foliage production was relatively similar between the two forests. Our results of aboveground biomass increments and fine-root production suggest a possible trade-off between carbon allocation to fine-roots versus wood growth (as it has been reported by other authors), as opposed to the most commonly assumed trade-off between total above- and below-ground production. Despite these differences among forests in terms of carbon allocation components, the leaf area index showed differences between forests like total NPP, suggesting that the leaf area index is more indicative of total NPP than carbon allocation. In Chapter 4, I evaluated the spatial and temporal variation of carbon allocation components and forest dynamics of Amazon forests as well as their responses to climatic fluctuations. I evaluated the intra- and inter-annual variation of carbon allocation components and forest dynamics during the period 2004−2012 in five forests on different soils (clay, clay-loam, sandy-clay-loam, sandy-loam and loamy-sand), but growing under the same local precipitation regime in north-western Amazonia (Colombia). We were interested in examining if these forests respond similarly to rainfall fluctuations as many models predict, considering the following questions: (i) Is there a correlation in carbon allocation components and forest dynamics with precipitation? (ii) Is there a correlation among forests? (iii) Are temporal responses in leaf area index (LAI) indicative of variations of above-ground production or a reflection of changes in carbon allocation patterns among forests?. Overall, the correlation of above- and below-ground carbon allocation components with rainfall suggests that soils play an important role in the spatial and temporal differences of responses of these forests to rainfall fluctuations. On the one hand, most forests showed that the above-ground components are susceptible to rainfall fluctuations; however, there was a forest on loamy-sand that only showed a correlation with the below-ground component (fine-roots). On the other hand, despite the fact that north-western Amazonia is considered without a conspicuous dry season (defined as <100 mm month−1), litterfall and fine-root mass showed high seasonality and variability, particularly marked during the drought of 2005. Additionally, forests of the loam-soil group showed that litterfall respond to time-lags in rainfall as well as and the fine-root mass of the loamy-sand forest. With regard to forest dynamics, only the mortality rate of the loamy-sand forest was significantly correlated with rainfall (77%). The observed inter-annual variability of stem and biomass increments of individuals highlighted the importance of the mortality in the above-ground biomass increment. However, mortality rates and death type proportion did not show clear trends related to droughts. Interestingly, litterfall, above-ground biomass increment and recruitment rates of forests showed high correlation among forests, particularly within the loam-soil forests group. Nonetheless, LAI measured in the most contrasting forests (clay-soil and loamysand) was poorly correlated with rainfall but highly correlated between forests; LAI did not reflect the differences in the carbon allocation components, and their response to rainfall on these forests. Finally, the forests studied highlight that north-western Amazon forests are also susceptible to climate fluctuations, contrary to what has been proposed previously due to their lack of a pronounced dry season.

Climatic analysis methodology of vernacular architecture

Vernacular architecture has demonstrated its perfect environmental adaptation through its empirical development and improvement by generations of user-builders. Nowadays, the sustainability of vernacular architecture is the aim of some research projects in which the same method should be applied in order to be comparable. Hence, we propose a research method putting together various steps. Through the analysis of geographical, lithology, economic, cultural and social influence as well as materials and constructive systems, vernacular architecture is analyzed. But, all this information is put together with the natural landscape (topography and vegetation) and the climatic data (temperature, winds, rain and sun exposure). In addition, the use of bioclimatic charts, such as Olgyay or Givoni’s, revealed the necessities and strategies in urban and building design. They are satisfied in the vernacular architecture by the application of different energy conservation mechanisms, some of them are shown by different examples in this paper.

Bio-climatic Design Handbook: designing public space to reach urban sustainability.

The "Bio-climatic Design Handbook: guidelines for the development of planning regulations" is a tool for urban planning and design professionals planning for the construction of public space taking into account bioclimatic and environmental standards. Based on environmental conditions assessment, urban design guidelines are given. These take into account various scales; from the territory to the microclimatic reality. From these general keys for the design of public space the handbook performs recommendations on specific case studies. The application of bioclimatic techniques in urban design promotes comfort in the public space and the respect for the existing environment, while it influences the energy consumption of buildings that conform this open space. The tool was developed in the context of BIOURB project, where Spain and Portugal cooperate writing this bilingual handbook. The case studies are located in this cross-border region.

Unravelling past flash flood activity in a forested mountain catchment of the Spanish Central System

Flash floods represent one of the most common natural hazards in mountain catchments, and are frequent in Mediterranean environments. As a result of the widespread lack of reliable data on past events, the understanding of their spatio-temporal occurrence and their climatic triggers remains rather limited. Here, we present a dendrogeomorphic reconstruction of past flash flood activity in the Arroyo de los Puentes stream (Sierra de Guadarrama, Spanish Central System). We analyze a total of 287 increment cores from 178 disturbed Scots pine trees (Pinus sylvestris L.) which yielded indications on 212 growth disturbances related to past flash flood impact. In combination with local archives, meteorological data, annual forest management records and highly-resolved terrestrial data (i.e., LiDAR data and aerial imagery), the dendrogeomorphic time series allowed dating 25 flash floods over the last three centuries, with a major event leaving an intense geomorphic footprint throughout the catchment in 1936. The analysis of meteorological records suggests that the rainfall thresholds of flash floods vary with the seasonality of events. Dated flash floods in the 20th century were primarily related with synoptic troughs owing to the arrival of air masses from north and west on the Iberian Peninsula during negative indices of the North Atlantic Oscillation. The results of this study contribute considerably to a better understanding of hazards related with hydrogeomorphic processes in central Spain in general and in the Sierra de Guadarrama National Park in particular.