Aridity Gradients Shape Intraspecific Variability of Morphological Traits in Native Ceratonia siliqua L. of Morocco.

The carob tree (Ceratonia siliqua L.) is a significant fruit tree in the Mediterranean region with cultural, biological, and ecological importance. Despite its importance, intraspecific trait variability (ITV) in carob trees has been largely overlooked in previous studies. Understanding ITV and its relationship with environmental conditions is crucial for conservation and breeding programs. In this study, we investigated the variability of carob pod and seed-related traits across different ecological scales in 25 studied populations in Morocco. Significant differences in morphological traits were observed between carob populations at various ecological levels, and pod-related traits exhibited greater variability than seed traits. Correlation analysis revealed strong associations between carob morphological traits and environmental conditions, with altitude and aridity index playing an influential role. The aridity gradient was strongly related to changes in pod size, seed number, and size, as well as seed yield. Our findings highlight an important ITV reaching 45% at the intra-population level, 36.5% at the inter-geographic level, and 30% at the inter-population level. Overall, this study contributes valuable insights into the ecology and adaptation of carob trees, emphasizing the importance of considering intraspecific variability when studying this remarkable species. This knowledge is critical for addressing the challenges posed by climate change and human activities on the long-term survival and ecological functioning of carob populations.

Quantifying Leaf Trait Covariations and Their Relationships with Plant Adaptation Strategies along an Aridity Gradient.

A trait-based approach is an effective way to quantify plant adaptation strategies in response to changing environments. Single trait variations have been well depicted before; however, multi-trait covariations and their roles in shaping plant adaptation strategies along aridity gradients remain unclear. The purpose of this study was to reveal multi-trait covariation characteristics, their controls and their relevance to plant adaptation strategies. Using eight relevant plant functional traits and multivariate statistical approaches, we found the following: (1) the eight studied traits show evident covariation characteristics and could be grouped into four functional dimensions linked to plant strategies, namely energy balance, resource acquisition, resource investment and water use efficiency; (2) leaf area (LA) together with traits related to the leaf economic spectrum, including leaf nitrogen content per area (Narea), leaf nitrogen per mass (Nmass) and leaf dry mass per area (LMA), covaried along the aridity gradient (represented by the moisture index, MI) and dominated the trait-environmental change axis; (3) together, climate, soil and family can explain 50.4% of trait covariations; thus, vegetation succession along the aridity gradient cannot be neglected in trait covariations. Our findings provide novel perspectives toward a better understanding of plant adaptations to arid conditions and serve as a reference for vegetation restoration and management programs in arid regions.

Resisting Aridification: Adaptation of Sap Conduction Performance in Moroccan Wild Olive Subspecies Distributed Over an Aridity Gradient.

In the current context of global change, the increasing frequency and the length of drought periods are testing the resistance capacities of plants of dry habitats. However, although the adaptation of plants to drought has been widely studied, the anatomical features of wood influencing the functional responses of plants to drought are still lacking at the intraspecific level, especially for species with a wide geographical distribution. As a result, we have studied the variation of wood anatomical traits related to sap conduction (i.e., vessel surface area, vessel density, and number of vessels joined by radial file) in two wild olive subspecies distributed in Morocco (i.e., Olea europaea subsp. europaea. var. sylvestris and Olea europaea subsp. maroccana), in relation to various drought conditions. This functional study, based on wood trait measurements of 351 samples from 130 trees and 13 populations, explores potential sap conduction in relation to environmental parameters and as a result, strategies to resist water stress. We found that (1) branch diameter (BD) captured 78% of total wood trait variation, (2) vessel size (SVS) expressed 32% of intraspecific variation according to cambium age, and (3) the positive relationship between SVS and BD could be explained by climate type, vegetation cover changes, and therefore available water resources. Taking into consideration the diameter of the branch as the main factor of anatomical variation, established reaction norms (linear models) at the intrapopulation scale of vessel lumen area according to aridity show for the first time how the functioning of the cambium modulates and controls sap conduction, according to aridity and thus available water resources. They pinpoint the risks incurred by the wild olive tree in the perspective of a dramatic increase in aridity, in particular, the inability of the cambium to produce large enough vessels to efficiently transport sap and irrigate the leaves. Finally, this study opens new and interesting avenues for studying at a Mediterranean scale, the resistance and the vulnerability of wild forms and cultivated varieties of olive to heterogeneous and changing environmental conditions.

Trait-based plant ecology a flawed tool in climate studies? The leaf traits of wild olive that pattern with climate are not those routinely measured.

Climate-related studies have generally focussed upon physiologically well-defined 'mechanistic' traits rather than 'functional' ones relating indirectly to resource capture. Nevertheless, field responses to climate are likely to typically include both 'mechanistic' specialization to climatic extremes and 'functional' strategies that optimize resource acquisition during less climatically-severe periods. Here, this hypothesis was tested. Seventeen traits (six 'functional', six 'mechanistic' and five 'intermediate') were measured from 19 populations of oleaster (wild olive) along a climatic gradient in Morocco. Principal components analysis of the trait dataset identified size and the 'worldwide leaf economics spectrum' as PCA axes 1 and 2. However, contrary to our prediction, these axes, and commonly-measured 'functional' traits, were little correlated with climate. Instead, PCA 3, perhaps relating to water-use and succulence, together stomatal density, specific leaf water content and leaf shape, patterned with altitude, aridity, rainfall and temperature. We concluded that, at least for slow-growing species, such as oleaster, 'mechanistic' traits are key to identifying mechanisms of climatic restriction. Meaningful collaboration between 'mechanistic' and 'functional' disciplines provides the best way of improving our understanding of the global impacts of climate change on species distribution and performance.