Rafael Calama

To grow or to seed: ecotypic variation in reproductive allocation and cone production by young female Aleppo pine (Pinus halepensis, Pinaceae)

Age and size at the first reproduction and the reproductive allocation of plants are linked to different life history strategies. Aleppo pine only reproduces through seed, and, as such, early female reproduction confers high fitness in its infertile highly fire-prone habitats along the Mediterranean coast because life expectancy is short. We investigated the extent of ecotypic differentiation in female reproductive allocation and examined the relation between early female reproduction and vegetative growth. In a common-garden experiment, the threshold age and size at first female reproduction and female reproductive allocation at age seven differed significantly among Aleppo pine provenances of ecologically distinct origin. Significant correlations among reproductive features of the provenances and the ecological traits of origin were found using different analytical tools. In nonlinear models of cone counts vs. stem volume, medium-sized trees (not the largest trees) produced the highest cone yield, confirming that, at the individual level, early female reproduction is incompatible with fast vegetative growth. The contribution of founder effects and adaptation to contrasting fire regimes may be confounding factors. But considering all traits analyzed, the geographical patterns of resource allocation by Aleppo pine suggest ecotypic specialization for either resource-poor (favoring early reproduction) or resource-rich (favoring vegetative growth) habitats.

Cone and seed production from stone pine (Pinus pinea L.) stands in Central Range (Spain)

Natural regeneration of stone pine (Pinus pinea L.) stands in the Central Range of Spain can be difficult to attain. The success of this regeneration is affected by factors such as the total amount of seed available, the short dispersal ability of the pinyon, the conditions required for germination, specific problems associated with the establishment, early survival of the seedlings due to severe summer heat and drought, competition for water and mineral resources and grazing damage. This study focuses on seed availability. The amount of available seeds depends on the number and size of the cones and the number of viable pinyons within the cones. In stone pine stands, both variables show great year to year variability. Both cone and seed production, for a given year, are also conditioned by the vigour and health of the tree, its size, the condition and attributes of the stand and the loss of seed through pests or predation. In this study, the main factors which influence cone and pinyon production are identified and a multivariate model to predict annual cone and viable seed production is developed. To consider the correlation among observations coming from the same tree, stand or year, random components are included in the model. The multivariate random structure allows for future calibration of the model for a given year from a small additional sample of observations. It is important to know the total amount of viable seed produced in a stand for a given year since regeneration cuttings for Stone pine should be concentrated in high crop years.

Thinning intensity and growth response in SW-European Scots pine stands

The effect of different thinning intensities on growth and yield was studied in Pinus sylvestris L. stands at the south-western limit of its distribution area (Central Spain), using five long-term thinning trials. Data were analysed collectively considering several factors (trial, block, plot and period) as random effects. Total volume and volume increment decreased with thinning intensity, this loss being more significant in the case of moderate and heavy thinning. No difference was found among treatments for total basal area or the increment in basal area. The results revealed an optimum basal area (Assmann’s definition) between 85 and 100% of the basal area in unthinned plots. Volume growth loss associated with heavy thinnings (reduction of 18% in volume increment) was smaller than that reported in Central and Northern European regions (greater than 25%). Height increment was not influenced by thinning, whereas dominant and quadratic mean diameter increments increased with the thinning intensity. The response of diameter growth to thinning was greater at younger ages (less than 50 y) and in medium-sized trees.RésuméL’effet de différentes intensités d’éclaircie sur la croissance et la production de peuplements de Pinus sylvestris L. a été étudié à la limite sud-ouest de sa zone de répartition (centre de l’Espagne), à partir de cinq essais d’éclaircie suivis à long terme. Les données ont été analysées collectivement en prenant en compte différents facteurs (essai, bloc, placeau et période) comme facteurs aléatoires. Le volume total et l’accroissement en volume diminuaient avec l’intensité d’éclaircie, cette perte devenant plus significative dans le cas d’éclaircies modérées et fortes. Aucune différence n’a été trouvée entre traitements pour la surface terrière totale ni pour sa vitesse d’accroissement. Les résultats révèlent un optimum de surface terrière (selon la définition d’Assmann) compris entre 85–100 % de la surface terrière des placeaux non éclaircis. Les pertes de croissance en volume associées aux éclaircies fortes (réduction de 18 %) étaient plus faibles que celles qui sont rapportées pour les régions du Centre et du Nord de l’Europe (supérieures à 25 %). L’accroissement en hauteur n’a pas été influencé par l’éclaircie alors que l’accroissement quadratique moyen en diamètre augmentait avec l’intensité d’éclaircie. La réponse en termes d’accroissement en diamètre à l’éclaircie a été plus forte pour les jeunes arbres (moins de 50 ans) et les arbres de taille moyenne.

Difference in cuticular transpiration and sclerophylly in juvenile and adult pine needles relates to the species-specific rates of development

Pinus species show remarkable ontogenetic differences in needle morphology (heterophylly) between juvenile and adult vegetative phases. This developmental shift may play an adaptative role in their success under diverse habitats. As a first step to know the functional differences between each vegetative phase, we compared water loss through the cuticles of juvenile and adult needles of 21-month-old nursery-grown seedlings of nine hard pine species. Cuticular transpiration (CT), calculated after complete stomatal closure, was obtained by leaf-drying curves, and was related to leaf, ontogenetic and climatic parameters. The rate of cuticular transpiration (RCT) between juvenile and adult needles differed across pine species, and in particular segregated the Mediterranean species Pinus canariensis and P. halepensis, from the Eurasian P. uncinata and introduced species P. radiata. For these species, RCT was always higher in juvenile needles. The different leaf and ontogenic parameters studied were correlated with the variation in RCT among the nine pine species. We discuss this relationship in the light of the species ecology. Besides their possible adaptive interpretation, these results suggest an underlying need to consider the ontogenetic heterophylly when assessing functional traits in hard pine seedlings, in particular those traits that govern water relations.

Variables influencing cork thickness in Spanish cork oak forests: A modelling approach

In this study, we evaluate the influence of different variables on cork thickness in cork oak forests. For this purpose, first we fitted a multilevel linear mixed model for predicting average cork thickness, and then identified the explanatory covariates by studying their possible correlation with random effects. The model for predicting average cork thickness is described as a stochastic process, where a fixed, deterministic model, explains the mean value, while unexplained residual variability is described and modelled by including random parameters acting at plot, tree, plot × cork harvest and residual within-tree levels, considering the spatial covariance structure between trees within the same plot. Calibration is carried out by using the best linear unbiased predictor (BLUP) theory. Different alternatives were tested to determine the optimum subsample size which was found to be appropriate at four trees. Finally, the model was applied and its performance in the estimation of cork production was tested and compared with the cork weight model traditionally used in Spain.RésuméDans cette étude, nous avons mesuré l’influence de diverses variables sur l’épaisseur du liège des forêts de chênes-lièges. Dans ce but nous avons d’abord appliqué un modèle linéaire mixte pour prédire l’épaisseur moyenne du liège, et on a alors identifié les co-variables explicatives pour expliquer leur possible corrélation avec des effets aléatoire. Le modèle prédisant l’épaisseur moyenne du liège peut être décrit comme un processus stochastique où un modèle fixe et déterministe explique la valeur moyenne, tandis qu’une variabilité résiduelle inexpliquée est décrite et modélisée par l’inclusion de paramètres aléatoires relevant de la parcelle, de l’arbre, de la récolte de liège par parcelle et aux niveaux résiduels des arbres prenant en compte la covariance de la structure spatiale entre les arbres d’une même parcelle. Le calibrage a été réalisé en employant la théorie BLUP (Best linear unbiased predictor ou Meilleur prédicteur linéaire non biaisé) On a essayé différentes options pour trouver la dimension optimale de l’échantillon et on a trouvé qu’il était opportun d’utiliser quatre arbres par parcelles. Finalement le modèle a été appliqué pour calculer la production de liège et a été comparé avec le poids de liège obtenu avec le modèle employé d’habitude en Espagne.

Carbon sequestration in Mediterranean pine forests

Management objectives and techniques in woodlands dominated by Quercus species have changed dramatically over the last few decades, especially in the case of coppices. The fact that these stands are of little economic importance today is in part compensated by their value, for example, as carbon sinks. The identification of this important role of Quercus woodlands allows us to contemplate the development of specific silviculture and to promote investment in the stands. It is also important, in the context of global change, to analyze the dynamics involved in the transformation of Quercus woodlands from carbon sinks to emitters.

The role of developmental stage in frost tolerance of Pinus pinea L. seedlings and saplings

ContextAlthough drought is generally considered the main environmental constraint in Mediterranean environments, the ability to acclimate to and tolerate frost in early developmental stages can be a determinant for seedling survival of many Mediterranean tree species like stone pine (Pinus pinea L.).AimsThe aim of this study was to assess the impact of the developmental stage of naturally regenerated stone pine individuals on tolerance to low temperature (LT) from summer to late autumn and in spring, at a highly continental site in central Spain. Specifically, we tested to what extent the differences in tolerance are related to shoot heteroblasty.MethodsWe assessed LT tolerance of needles from individuals at three age classes (class C1: seedlings, class C2: 4- to 8-year-old saplings and class C3: >9-year-old saplings) over nine dates from summer to spring.ResultsLT tolerance displayed severe seasonal trends and differed between age classes. It usually increased with sapling age. Such differences were tightly related to heteroblasty of the shoots. Our results point to a higher LT tolerance associated with larger leaf dry mass per unit area (LMA) values. No impact of late frosts on shoot growth rates was detected during this study.ConclusionsDevelopmental changes during early plant growth seem to play a role in frost tolerance of stone pine seedlings, a finding which furthers our understanding of regeneration dynamics in this species in areas with continental influence.

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

Abstract• ContextPinus 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.• AimsDetermining 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.• MethodsAn 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.• ResultsDecreasing 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.• ConclusionsSurvival 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.

A silviculture-oriented spatio-temporal model for germination in Pinus pinea L. in the Spanish Northern Plateau based on a direct seeding experiment

Natural regeneration in Pinus pinea stands commonly fails throughout the Spanish Northern Plateau under current intensive regeneration treatments. As a result, extensive direct seeding is commonly conducted to guarantee regeneration occurrence. In a period of rationalization of the resources devoted to forest management, this kind of techniques may become unaffordable. Given that the climatic and stand factors driving germination remain unknown, tools are required to understand the process and temper the use of direct seeding. In this study, the spatio-temporal pattern of germination of P. pinea was modelled with those purposes. The resulting findings will allow us to (1) determine the main ecological variables involved in germination in the species and (2) infer adequate silvicultural alternatives. The modelling approach focuses on covariates which are readily available to forest managers. A two-step nonlinear mixed model was fitted to predict germination occurrence and abundance in P. pinea under varying climatic, environmental and stand conditions, based on a germination data set covering a 5-year period. The results obtained reveal that the process is primarily driven by climate variables. Favourable conditions for germination commonly occur in fall although the optimum window is often narrow and may not occur at all in some years. At spatial level, it would appear that germination is facilitated by high stand densities, suggesting that current felling intensity should be reduced. In accordance with other studies on P. pinea dispersal, it seems that denser stands during the regeneration period will reduce the present dependence on direct seeding.

Defining the optimal regeneration niche for Pinus pinea L. through physiology-based models for seedling survival and carbon assimilation

Key messageSeedling survival inPinus pineais controlled by both water status and photosynthetic performance. Optimal regeneration niche for the species is found at mid-shaded sites.AbstractSummer survival has been identified as the main bottleneck preventing natural regeneration in Mediterranean forests, although the physiological processes resulting in seedling mortality are not sufficiently known. In the present work, the effect of water status and photosynthetic performance on seedling survival and regeneration niche in a Mediterranean pine (Pinus pinea L.) was analyzed by means of a modeling approach. Midday water potential was modeled as a nonlinear function of environmental factors, and this model was coupled with an existing model for carbon assimilation. A model for seedling survival was then constructed using lifetime analysis techniques, including predicted values of daily and cumulative net assimilation and probability for critical midday water potentials as predictors. The model was applied over a wide range of irradiance environments in order to identify the optimal regeneration niche for the species. Results indicate that midday water potential for P. Pinea seedlings is affected by relative soil water content, leaf temperature, and irradiance, with younger seedlings being more likely to reach critical values. Seedling survival in P. pinea is controlled by both water status and photosynthetic performance, with mortality being triggered by the joint occurrence of low water potentials and negative assimilation rates, although photoassimilates stored during the spring season increase survival in older seedlings. Simulations indicated that seedling survival is optimized in shaded environments, while carbon assimilation reaches maximum values on more open sites, thus the optimal regeneration niche for the species is found on mid-shaded locations.