Signatures of natural selection in a foundation tree along Mediterranean climatic gradients

Temperature and precipitation regimes are rapidly changing, resulting in forest dieback and extinction events, particularly in Mediterranean-type climates (MTC). Forest management that enhance forests’ resilience is urgently required, however adaptation to climates in heterogeneous landscapes with multiple selection pressures is complex. For widespread trees in MTC we hypothesized that: patterns of local adaptation are associated with climate; precipitation is a stronger factor of adaptation than temperature; functionally related genes show similar signatures of adaptation; and adaptive variants are independently sorting across the landscape. We sampled 28 populations across the geographic distribution of Eucalyptus marginata (jarrah), in South-west Western Australia, and obtained 13,534 independent single nucleotide polymorphic (SNP) markers across the genome. Three genotype-association analyses that employ different ways of correcting population structure were used to identify putatively adapted SNPs associated with independent climate variables. While overall levels of population differentiation were low (FST = 0.04), environmental association analyses found a total of 2336 unique SNPs associated with temperature and precipitation variables, with 1440 SNPs annotated to genic regions. Considerable allelic turnover was identified for SNPs associated with temperature seasonality and mean precipitation of the warmest quarter, suggesting that both temperature and precipitation are important factors in adaptation. SNPs with similar gene functions had analogous allelic turnover along climate gradients, while SNPs among temperature and precipitation variables had uncorrelated patterns of adaptation. These contrasting patterns provide evidence that there may be standing genomic variation adapted to current climate gradients, providing the basis for adaptive management strategies to bolster forest resilience in the future

Signatures of natural selection in a foundation tree along Mediterranean climatic gradients

Temperature and precipitation regimes are rapidly changing, resulting in forest dieback and extinction events, particularly in Mediterranean-type climates (MTC). Forest management that enhance forests’ resilience is urgently required, however adaptation to climates in heterogeneous landscapes with multiple selection pressures is complex. For widespread trees in MTC we hypothesized that: patterns of local adaptation are associated with climate; precipitation is a stronger factor of adaptation than temperature; functionally related genes show similar signatures of adaptation; and adaptive variants are independently sorting across the landscape. We sampled 28 populations across the geographic distribution of Eucalyptus marginata (jarrah), in South-west Western Australia, and obtained 13,534 independent single nucleotide polymorphic (SNP) markers across the genome. Three genotype-association analyses that employ different ways of correcting population structure were used to identify putatively adapted SNPs associated with independent climate variables. While overall levels of population differentiation were low (FST = 0.04), environmental association analyses found a total of 2336 unique SNPs associated with temperature and precipitation variables, with 1440 SNPs annotated to genic regions. Considerable allelic turnover was identified for SNPs associated with temperature seasonality and mean precipitation of the warmest quarter, suggesting that both temperature and precipitation are important factors in adaptation. SNPs with similar gene functions had analogous allelic turnover along climate gradients, while SNPs among temperature and precipitation variables had uncorrelated patterns of adaptation. These contrasting patterns provide evidence that there may be standing genomic variation adapted to current climate gradients, providing the basis for adaptive management strategies to bolster forest resilience in the future

Biomass production in intensive plantations of Eucalyptus species in Uruguay

La producción de biomasa con especies de eucaliptos muestra un alto potencial, teniendo en cuenta los altos niveles de crecimiento que tienen estas especies en los diferentes suelos de prioridad forestal de Uruguay. La biomasa de estas especies (en particular la madera) presentan características interesantes desde el punto de vista tecnológico teniendo en cuenta la densidad básica, la homogeneidad de la composición química, el bajo contenido de azufre, cenizas y nitrógeno y un poder calórico relativamente alto. Estas características determinan que las plantaciones intensivas de estas especies puedan ser una alternativa interesante en sistemas de producción cuyo objetivo sea la obtención de distintos tipos de biocombustibles. Los cultivos de altas densidades y corta rotación tienen particularidades que merecen especial atención desde el punto de vista productivo, tecnológico y ambiental teniendo en cuenta la falta de información que hay sobre los mismos en los suelos forestales de Uruguay. En función de estas necesidades se instalaron en la primavera de los 2010 dos ensayos a campo en dos localizaciones (Tacuarembó-NE, y Paysandú-SW), con tipos de suelos representativos de los suelos predominantes en las regiones litoral y norte del país. Fueron evaluadas tres especies de eucaliptus (Eucalyptus benthamii, E. dunnii y E. grandis) establecidas con cuatro densidades de plantación (2220, 3330, 4440 y 6660 árboles por hectárea) en un diseño de parcelas divididas con tres repeticiones. El Capítulo 1 presenta un contexto general de la matriz energética en Uruguay y la conceptualización de la biomasa forestal y, en particular, de los cultivos energéticos; además de su posible adaptación a las condiciones del país y su potencial como materia prima para la obtención de combustibles. En función de ese contexto se plantean el objetivo general y los específicos los cuales se relacionan directamente con los capítulos de la presente tesis. En el Capítulo 2 se justifica la elección de las especies de eucaliptos y se analiza la distribución potencial de E. grandis y E. dunnii en los distintos tipos de suelos de prioridad forestal del país. E. benthamii no fue incluida en este primer análisis ya que no se contaban con datos suficientes de inventario por ser una especie de uso relativamente reciente por las empresas productoras de pasta de celulosa. Los resultados permitieron identificar los parámetros de suelo, topografía y clima que mayor asociación muestran con el crecimiento de estas especies y, por tanto, generar una cartografía de las regiones con mayor potencial para el crecimiento de ambas especies. Los modelos de predicción de hábitat ajustados mostraron un grado elevado de precisión, y también permiten visualizar la distribución a futuro de estas especies en función de diferentes escenarios de cambio climático. El Capítulo 3 analiza la evolución de la sobrevivencia y su relación con las especies y densidades de plantación en los primeros 5 años de crecimiento. La evolución de la sobrevivencia también es analizada en función de las variaciones de la temperatura y precipitaciones ocurridas durante el período de evaluación en ambos sitios, así como de las condiciones de preparación previo e inmediatamente posterior a la plantación. Fueron analizados los efectos de la competencia entre individuos y sus efectos sobre el volumen individual y por hectárea. La evolución de las curvas de crecimiento indica el momento óptimo de cosecha para cada caso. En el Capítulo 4 se presenta la dinámica de la densidad de la madera a lo largo de los ciclos de cultivo, la cual muestra un comportamiento diferente en ambas localizaciones para las especies y densidades de plantación en las diferentes edades evaluadas. El análisis del peso individual mostró un comportamiento similar al del volumen en cada una de las densidades de plantación. El análisis del efecto de la competencia en las diferentes densidades de plantación se manifiesta de forma similar al del volumen individual determinando diferencias en la productividad por hectárea de las especies en cada uno de los espaciamientos. Con estos resultados es posible identificar las especies y densidades de plantación con una productividad más elevada, y el momento óptimo de cosecha en cada caso. Al mismo tiempo, se observó la relación del espaciamiento y del turno de cosecha. Fueron identificadas las variables que permiten estimar con mayor precisión tanto del volumen como el peso individual para cada localidad, especie y densidad de plantación. En el capítulo 5 se analizan los siguientes parámetros energéticos: poder calórico superior, densidad de la madera, densidad energética y rendimiento energético por hectárea y por año. Fueron analizados los efectos de la localidad, la edad, la especie y la densidad a partir de los parámetros descritos, a los efectos de identificar la combinación de factores que maximiza la producción de energía por unidad de superficie y de tiempo. A su vez, fueron analizadas las relaciones entre la densidad de la madera y el poder calórico, y el peso relativo de estas en la densidad energética. Finalmente fue estimada la superficie plantada necesaria para abastecer una planta de generación de energía eléctrica de 10 MWh en cada localidad estudiada. En el Capítulo 6 se evalúan los niveles de producción de biomasa considerando todas las fracciones aéreas del árbol (madera, corteza, ramas y hojas), para cada especie y densidad de plantación. Se estudian los efectos de la competencia entre árboles sobre la proporción del peso de cada fracción con respecto a la biomasa total. Se analizan las concentraciones de nutrientes (P, Ca, Mg, K y N) de las fracciones madera, corteza y hojas en cada especie y densidad de plantación. Los contenidos de nutrientes entre fracciones muestran la relación de proporciones entre las mismas debido a las diferencias en la actividad fisiológica de cada una. El estudio de las concentraciones de nutrientes en cada fracción también permite visualizar la diferente distribución de estos en los diferentes componentes de la biomasa y las implicancias de extracción considerando las fracciones madera versus el árbol entero. La estimación de los kg de nutrientes extraídos para cada especie y densidad de plantación dan una idea clara de la fertilización que sería necesario reponer en el suelo para conseguir la sostenibilidad de este tipo de sistemas de producción. La cantidad de nutriente extraído versus la cantidad de biomasa producida da una idea de la eficiencia con que son utilizados los diferentes nutrientes en cada combinación especie, densidad de plantación y sitio. El balance del stock de cationes en el suelo versus las cantidades de nutrientes extraídos determina el número de rotaciones en los que sería posible mantener los actuales niveles de crecimiento asumiendo turnos de cosecha próximo a los seis años. Esto, a su vez, muestra cuál de los tres cationes evaluados (Ca, Mg y K) se tornarían restrictivos para el crecimiento en el corto y mediano plazo. En el Capítulo 7 se presenta la discusión general de la tesis y señalando nuevas interrogantes que surgen en este tipo de cultivos energéticos en los turnos siguientes a la primera cosecha teniendo en cuenta las particularidades de estos en esas etapas de crecimiento. Finalmente, en el Capítulo 8 se puntualizan las principales conclusiones obtenidas.Production of biomass from eucalyptus species shows a high potential taking into account their high levels of growth in the different forest priority soils in Uruguay. The biomass from these species (particularly wood) presents interesting characteristics from the technological point of view considering basic density, homogeneity of the chemical composition, the low content of sulphur, ash and nitrogen and a relatively high caloric power. These characteristics determine that these species could be an interesting alternative in production systems whose production objective is to obtain different types of biofuels. High density and short rotation plantations have particularities that need special attention from the productive, technological and environmental points of view, taking into account there is a lack of information on them in Uruguayan forest soils. Based on these needs, two field trials were installed in the spring 2010, at two sites with soil types which were representative of the predominant ones in the coastal and northern regions of the country. Three species of eucalyptus (Eucalyptus benthamii, E. dunnii and E. grandis) were evaluated in four planting densities (2220, 3330, 4440 and 6660 trees per hectare) in a split plot design with three replications. Chapter 1 presents a general context of the energy matrix in Uruguay and the conceptualization of forest biomass and, in particular, of energy crops as well as their possible adaptation to the country's conditions as well as their potential as raw material for fuels. Depending on this context, the general objective and the specific ones are established, which are directly related to the chapters of this thesis. In Chapter 2 the choice of eucalyptus species is justified and the distribution potential of E. grandis and E. dunnii is analysed in the different types of forest priority soils in the country. E. benthamii was not included in this first analysis since there was not enough inventory data as it is a relatively recent species used by pulp producers. Results allowed to identify the parameters of soil, topography and climate that show a greater association with their growth and therefore a mapping of regions with greater potential for the growth of both species. The models for adjusted habitat prediction showed a high degree of precision and also allowed to visualize a future distribution of these species depending on different climate change scenarios. Chapter 3 analyses survival evolution and its relationship with species and planting densities in the first 5 years of growth. The evolution of survival is also analysed according to the variations in temperature and rainfall during the evaluation period in both sites and also from the point of view of soil preparation conditions prior to and immediately after planting. Competition effects between individuals and their effects on individual volume and per hectare were analysed as well. The evolution of the growth curves indicates the optimum harvest time in each case. In Chapter 4, dynamics of wood density are presented. A different behaviour is shown on both sites for the species and planting densities at the different evaluated ages. The analysis of the individual weight showed a similar behaviour to the volume in each of the planting densities. The analysis of the competition effect on the different densities of plantation manifests in a similar way to the individual volume, determining differences in the species productivity per hectare in each of the spacings. Based on these results it is possible to identify the species and plantation densities with higher productivity levels as well as the optimum harvest time in each case. At the same time, a relationship between spacing and harvest time was observed. The variables that allow to estimate with greater precision both volume and individual weight for each species site and density of plantation were identified. Chapter 5 discusses the following energy parameters: higher caloric power, wood density, energy density and energy yield per hectare and per year. The effects of site, age, species and density were analysed with each of these parameters in order to identify the combination of factors that maximizes energy production per area and time. In turn, relationships between wood density and caloric power and their relative weight in determining energy density were analysed. Finally, it was estimated the planted área needed to supply a 10 MWh power generation plant for each site. In Chapter 6, levels of biomass production are evaluated considering all the tree aerial fractions (wood, bark, branches and leaves) for each species as well as plantation density. It has also studied the effects of competition among trees with the proportion of each fraction weight in respect to the total biomass. The nutrient concentrations (P, Ca, Mg, K and N) are analysed in wood, bark and leaf fractions in each species and planting density. The nutrient contents in fractions show the proportion relationships among them due to the differences in the physiological activity of each one. The study of nutrient concentrations in each fraction also allows to visualize the different distribution pattern of these in the different components of the biomass and the extraction implications considering wood versus whole tree fractions. The estimation of kg of nutrients extracted for each species and plantation density gives a clear idea of mineral magnitudes that would be necessary to replenish the soil in order to achieve the sustainability of this type of production system. The amount of nutrient extracted versus the amount of biomass produced provides an idea of the efficiency with which different nutrients are used in each species combination, planting density and site. Balance of cations stock in the soil versus the amounts of extracting nutrients determines the number of rotations in which it would be possible to maintain current growth levels, assuming turns close to six years. This, in turn, shows which of the three evaluated cations (Ca, Mg and K) would become restrictive for growth in the short and medium term. Chapter 7 presents the general discussion of the thesis and points out new questions that arise in this type of energy crops in the turns following the first harvest, considering their particularities in those stages of growth. Finally, in Chapter 8 the main conclusions obtained are highlighted

Towards greater accuracy in individual-tree mortality regression

Background mortality is an essential component of any forest growth and yield model. Forecasts of mortality contribute largely to the variability and accuracy of model predictions at the tree, stand and forest level. In the present study, I implement and evaluate state-of-the-art techniques to increase the accuracy of individual tree mortality models, similar to those used in many of the current variants of the Forest Vegetation Simulator, using data from North Idaho and Montana. The first technique addresses methods to correct for bias induced by measurement error typically present in competition variables. The second implements survival regression and evaluates its performance against the traditional logistic regression approach. I selected the regression calibration (RC) algorithm as a good candidate for addressing the measurement error problem. Two logistic regression models for each species were fitted, one ignoring the measurement error, which is the “naïve” approach, and the other applying RC. The models fitted with RC outperformed the naïve models in terms of discrimination when the competition variable was found to be statistically significant. The effect of RC was more obvious where measurement error variance was large and for more shade-intolerant species. The process of model fitting and variable selection revealed that past emphasis on DBH as a predictor variable for mortality, while producing models with strong metrics of fit, may make models less generalizable. The evaluation of the error variance estimator developed by Stage and Wykoff (1998), and core to the implementation of RC, in different spatial patterns and diameter distributions, revealed that the Stage and Wykoff estimate notably overestimated the true variance in all simulated stands, but those that are clustered. Results show a systematic bias even when all the assumptions made by the authors are guaranteed. I argue that this is the result of the Poisson-based estimate ignoring the overlapping area of potential plots around a tree. Effects, especially in the application phase, of the variance estimate justify suggested future efforts of improving the accuracy of the variance estimate. The second technique implemented and evaluated is a survival regression model that accounts for the time dependent nature of variables, such as diameter and competition variables, and the interval-censored nature of data collected from remeasured plots. The performance of the model is compared with the traditional logistic regression model as a tool to predict individual tree mortality. Validation of both approaches shows that the survival regression approach discriminates better between dead and alive trees for all species. In conclusion, I showed that the proposed techniques do increase the accuracy of individual tree mortality models, and are a promising first step towards the next generation of background mortality models. I have also identified the next steps to undertake in order to advance mortality models further

Genetic variation and resilience to climate change in Mediterranean-type trees

Climate models predict that temperature means will continuously increase globally, and that heatwaves and drought periods will become more frequent and intense, particularly in Mediterranean-type climates. The Southwest Western Australia (SWWA) Biodiversity Hotspot has extensive forest environments that have been subject to heatwaves and drought-induced forest mortality in recent years, impacting forest carbon sequestration and local ecological structure. Although, species may persist through enhanced physiological tolerance, phenotypic plasticity and/or genetic adaptation. Genetic variation is critical for ecological adaptive capacity - the potential and ability to adjust to, and persist through, external factors - and consequently, the evolutionary potential of the species. Evolution to a specific environment through natural selection results in patterns of local adaptation (when a local population experiences higher fitness compared to non-local counterparts). Local adaptation can be identified by either genome wide surveys that link genetic variants to climate variables or measuring plant traits indicative of plant performance and survival through reciprocal transplants in common environments. Exploring genetic adaptation patterns associated with physiological tolerance to climate can guide forest management approaches to enhance forests’ resilience to climate change, such as assisted gene migration. The genetic survey (Chapter 2) sampled natural jarrah populations and obtained 13,534 independent single nucleotide polymorphic (SNP) markers across the genome. Three genotype-association analyses were used to identify putatively adapted SNPs associated with independent climate variables. While overall levels of population differentiation were low (FST=0.04), environmental association analyses found a total of 2,336 unique SNPs associated with temperature and precipitation variables, with 1,440 SNPs annotated to genic regions. Considerable allelic turnover was identified for SNPs associated with temperature seasonality and mean precipitation of the warmest quarter, suggesting that both temperature and precipitation are important factors in adaptation. SNPs with similar gene functions, had analogous allelic turnover along climate gradients, while SNPs among temperature and precipitation variables had uncorrelated patterns of adaptation. These contrasting patterns provide evidence that there may be standing genomic variation adapted to current climate gradients, providing the basis for adaptive management strategies to bolster forest resilience in the future. The second experimental chapter (Chapter 3) explored seed germination response to temperature in jarrah and marri populations from wide-ranging climate origins, to estimate the thermal optima and constraints. Seeds from across the entire geographic distribution were collected from independent populations of each species. Patterns of germination observed differences between species on a thermal gradient plate (5-40°C) and provided a temperature range for explicit germination tests. Germination tests were carried out at five constant temperatures between 9 and 33°C. We discuss how the germination niche (1) differs between species, (2) varies among populations, and (3) relates to climatic origin. Temperature response of germination differed among species, specifically the optimum temperature for germination (jarrah – 23.4°C; marri - 31°C). Temperature response of germination also differed among populations within species and was related to the climate-origin only for marri. Lastly, for the third experimental chapter (Chapter 4), a reciprocal transplant common garden experiment was used to investigate variation in marri`s functional traits using hyperspectral data. Hyperspectral remote sensing has the potential to assess plant functional status rapidly and non-destructively across climatic gradients to support conservation and management strategies, such as assisted migration, for forests under climate change. This study explored the variability of functional traits in marri to estimate patterns of local adaptation. Trees from natural populations spanning marri’s geographic distribution were grown in two common garden plantation sites with different climate settings. High-resolution field-based spectral measurements were collected from leaves of adult plants at both sites in two seasons (summer and autumn). Partial least squares regression analyses of full reflectance spectra highlighted differences among populations, sites, and seasons in spectral regions associated with photosynthetic pigments and water content, among other spectral traits, related to leaf condition and stress responses. Variation in these traits was further explored with analyses of spectral indices tailored to pigment and water absorptions. Analyses of spectral indices variation identified significant differences between populations, suggesting there is heritable variation in climatic tolerances, but stronger effects of season and site

Conservation Genetics for Management of Threatened Plant and Animal Species

This book focuses on conservation genetic (and genomic) papers that demonstrate applied outcomes that inform practical threatened species management. We cover a broad range of species and genetic approaches, but focus on how conservation genetic information is used to underpin management actions for species recovery. Through the exposition of a diversity of approaches, we aim to demonstrate to conservation managers and researchers how conservation genetics can inform on-ground species management

Models to support eucalyptus plantations management under a changing environment

Doutoramento em Engenharia Florestal e dos Recursos Naturais - Instituto Superior de AgronomiaThe objective of this work was the development of a tool to support eucalyptus stands management with the capacity of predicting forest development under different management options, thus providing forest managers with useful information in the form of variable outputs with interest for forest management. The tool uses the 3-PG model as the basis for growth predictions so that it can be used under a changing climate. The first thing to be done was the improvement of the crown ratio equation, taking advantage of the great amount of available data. With more precise equations, new values of biomass where estimated and a new set of aboveground biomass equations was developed. Biomass values are not only an important model output, but also a vital piece in the hybridization of the GLOBULUS growth and yield model and the 3-PG whole stand process based model. The linkage of the models resulted in a hybrid model with more detailed outputs that were further complemented with a diameter distribution model. The 3-PG fertility ratio is an important parameter of the model, but is estimated in a subjective way. The improvement with an equation that predicts it from soil characteristics is important not also because it is a flaw that is recurrently appointed to the model, but also because it will allow it to be sensible to fertilizationsFC

Opportunities in small population breeding in black cottonwood : or The little population that could

Small and structured populations are problematic for breeding due to low levels of genetic variability and increased levels of linkage disequilibrium (LD). The increased LD causes problems with utilization of common genomic tools and may lead into increased accruement of deleterious alleles through the intense selection and bottlenecks characteristic of breeding. In forest trees, maintaining a large breeding population to alleviate these issues is difficult due to large physical size of most trees. The aim of this thesis was to study and resolve some of the LD dependent issues of small populations and establish small populations as an option in breeding. We established the accuracy of LD-based estimation methods of recombination using data from European aspen, allowing confident use of these methods later in the project. The small black cottonwood breeding population contained enough genetic diversity to facilitate future adaptive selection to novel Swedish climate and light conditions, suggesting that small populations consisting of offspring with diverse parentage are feasible options for breeding in species with high outcrossing and recombination rates. We identified candidate genes that can be targeted for selection on phenology and growth using a genome wide association study (GWAS). We also show that GWAS is useful for identifying large effect alleles even in small populations and that efficient growth under novel conditions likely require different allele combinations than in native habitats. Finally, we quantified deleterious load and identified effects of the deleterious load on growth. Accounting for deleterious load allowed for more effective genomic selection and increased breeding cycle gain in breeding programs based on small populations

Ecotypic variation and plasticity of morphological and physiological traits of Eucalyptus loxophleba ssp. lissophloia along a climate gradient in south-west Australia

The adaptive capacity of plant species will be important for increasing their resilience in a changing climate. By 2070, a change in rainfall of +10 to -40 % of current mean annual rainfall and warming of between +2 to +5 ˚C in south-west Western Australia, is predicted. Plant species may cope with changing climatic characteristics through natural selection or phenotypic plasticity responses. Greater ecotypic variation in traits between stands of a single species may reduce its vulnerability in a changing climate as there is a greater selection of traits to increase fitness. Phenotypic plasticity has been highlighted as a mechanism to potentially enhance resistance and resilience in a changing climate in the short-term. Given that the Mediterranean region of the south-west of Western Australia is considered to be the most vulnerable to contraction of all Mediterranean systems under predicted climate change scenarios, the region is a priority area for research into climate resilience and adaptive ecotypic variation in plant species. This project examined differences in morphological and physiological traits between nine stands of the widespread species E. loxophleba ssp. lissophloia L.A.S. Johnson & K.D. Hill (Smooth-barked York Gum) across a climate gradient in south-western Australia. Morphological and physiological traits known to promote efficient water use and drought tolerance were compared across its natural range (450 km from W to E), with samples of the same provenances also grown in a common garden (plantation) which receives higher long-term average annual rainfall. Ten trees per natural stand and per provenance were selected for sampling. Traits measured were total leaf length, maximum leaf width, the ratio of leaf length to leaf width, area per leaf, dry mass, specific leaf area, leaf nitrogen, carbon and nitrogen isotope ratios, instantaneous water use efficiency, maximum photosynthetic and transpiration rate and wood density. The eastern end of the gradient tended to have lower long-term average rainfall and higher long-term average temperatures than the sites further west. However, in the twelve months prior to sampling (01/06/2010 to 31/05/2011) rainfall patterns were anomalous and sites to the west of the gradient received below average rainfall. This meant that the long-term rainfall gradient was actually inverted when considering annual rainfall in the twelve months prior to sampling. The natural stands along the climate gradient and the plantation were similar in their soil characteristics and biotic structure. Results showed significant differences (one-way ANOVA) between the nine natural stands along the climate gradient for all of the traits measured, with the exception of wood density. This showed the high ecotypic variation present within the widespread species in both foliar morphological and physiological traits measured. However, there was no single, major climatic variable which was the primary cause of variation of all the traits, as determined by linear regression analysis. Comparisons of the natural versus planted stands showed high phenotypic plasticity responses in the physiological traits to prevailing climatic conditions, which should allow the species to cope with climate changes of a limited scale over the short-term. Morphological leaf traits showed a more conservative response, although the high ecotypic variation between stands suggests there is variation to facilitate change if given sufficient time. In conclusion, some adaptive capacity of traits relating to drought tolerance and water conservation has been shown in a single species. Despite the ability to cope with a changing climate afforded by phenotypic responses and ecotypic variation within a species, the necessity of mitigating climate change remains paramount